Study Plan, Sierra Nevada Research Center
vs 02.06.07
1. Project Title
Regional climate variability and high-elevation forest response over the past 4000
years inferred from limber pine (Pinus flexilis) chronologies and forest stand
dynamics in the eastern Sierra Nevada and western Great Basin ranges.
Principal Investigator and Research Associates
Constance Millar (PI), with Robert Westfall, Diane Delany
Sierra Nevada Research Center (SNRC); and
John King
Lone Pine Research, Bozeman, MT
2. Problem Reference
Sierra Nevada Research Center RWUD
Problem 3: Climate and Landscape Change
Specific goals of Problem 3 include:
• Evaluate climate-induced changes in species composition, range distributions, forest structure and
function at century scales in the high Sierra Nevada
• Identify the role and magnitude of climate as an ecological architect relative to other landscape forces
• Provide meaningful and useful interpretations and applications to ecological restoration, conservation
and management
3. Literature Review and Background
Climate change and its effects on ecosystems are key current topics of ecological research and pressing
challenges for natural-resource management and conservation (Millar and Brubaker, in press; Millar,
2004; Millar & Woolfenden, 1999a, b; 2001). Despite the anthropogenic component to current climate
change, pre-historic “natural” climate change and associated ecological responses afford critical context
for understanding the range of natural variability and the dynamics of current forest systems. Climate
change has been little comprehended as a natural driver of ecological change within conservation and
management communities, and misassumptions abound of effects being simple, linear, and equilibrium
responses. Increased understanding of climate’s role as a “macro-disturbance” agent will enable better
estimates of future impacts on forests, and improved forest management planning in the Sierra Nevada
and elsewhere. Little detailed work exists, however, on the nature of forest response to climate and the
implications to management.
Recent advances in climate science emphasize the hierarchic nature of climate variability at different
scales, including interannual (e.g., El Niño/La Niña), multidecadal (e.g., Pacific Decadal Oscillation),
centennial –millennial (Bond Cycles), and multi-millennial (Milankovitch Cycles). These and other modes
act in concert to create climate on earth at any point in time, with complex interactions and nonequilibrium effects. Evidence is also increasing worldwide on the nature of ecological response to these
climate modes in the past, suggesting similarly complex ecological behavior, characterized by lags,
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threshold effects, episodic responses, and non-equilibrium conditions. Understanding these at regional
scales becomes essential for clarifying dynamics of local ecosystems and how 21st century global change
will affect native species.
The eastern Sierra Nevada, CA and southwestern Great Basin, NV have become important focal areas at
the international scale for research on paleoclimatology and paleoecology. The reasons for this are
several. Steep and complex climate gradients exist from the crest of the Sierra Nevada across the
eastern escarpment and into the western and central Great Basin. Complex climate patterns derive from
a combination of synoptic climatology patterns and highly localized storm tracks that are influenced by the
complex topography of the area. The combination of diverse topography, mountain ranges isolated by
intervening basins, and high climate variability contributes to complex ecological contexts, with each
mountain range having slightly different conditions. Thus, populations in each range become useful
subjects for intercomparison research, given their similarities and differences. Further, while there is
considerable climate variability, in general the climates are consistenty cold and semi-arid across the
eastern Sierra Nevada and western Basin ranges, affording an opportunity to compare finer scale climate
influences among sites.
Cumulatively this landscape also promotes conditions for preservation of many different indicators of past
climate and ecosystem conditions. These include preserved deadwood, marcrofossils from packrat
middens, and pollen, micro-fauna, and physical indicators in sediments from lake and playa bottoms. The
dominant conifer species of upland mountain forests in the region, limber pine and bristlecone pine (Pinus
flexilis and P. longaeva, respectively) are long-lived (longevities ca 2000 and 4000 yrs, respectively),
sensitive to climate, readily cross-datable by dendrochronological methods, and yield longer records than
many other species with informative climate records. In the upland regions, wildfire tends to have a small
and localized role as disturbance agent, and insect/pathogen activity is minimized by cold, arid conditions.
Thus, the effects of climate on ecological dynamics are clearer in this region than in warmer, wetter
climes where other disturbance factors confound climate analysis. Taken together, the many and
independent proxies for paleoenvironmental conditions in the region mean that intercomparisons are
possible, and corroborations are available to make historical inferences defensible. As a result, the
literature on many aspects of paleoenvironments is excellent, although many gaps remain in
understanding, making the region valuable for further research.
Focal Research Region and Study Areas
The focal research region is a rectangular area in central-eastern Sierra Nevada and southwestern Great
Basin. Within this region, the primary study area is described as follows (fig. 1): The western boundary
follows the Sierra Nevada crest. The northern boundary extends along the latitude of Markleeville, CA, ca
38.7°N; the southern boundary along the latitude of Mammoth Lks, CA , ca 37.6°N. The eastern
boundary parallels the Sierra Nevada crestline, along a line that extends from the eastern base of the
White/Inyo Mountains, CA/NV through the eastern base of the Wassuk Range, NV.
Additional outlying study sites within the general focal region include the northern Toiyabe Mountains and
the upper Owens Gorge (fig. 1). The elevation of the latter site is the lowest of all study areas, with a
base elevation of 2070 m.
Forest and Woodland Types
Rangeland shrubs and grasses dominate the lowest basins and the high alpine zones below and above
treeline, respectively. The lowest woodland type, mostly below our study areas, is Utah juniper
(Juniperus osteosperma), which occurs at 1300 – 2750 m, from the Sierra Nevada east through Nevada.
Utah juniper is dry-adapted but ranges broadly in elevation. Nested within its elevation range is singleneedle pinyon pine (Pinus monophylla), which associates with juniper throughout its elevation range of
1350 – 2665 m, but dominates mid-slopes. Pinyon/juniper and associated big sagebrush (Artemesia
tridentata)/bitterbrush (Purshia tridentate) shrub types are relatively constant through their elevation zone
in the general region.
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Above the pinyon/juniper woodland zone, higher elevation forest types vary widely over the focal region.
In the central eastern Sierra Nevada, conifer forests are abundant and diverse. Limber pine populations,
however, are highly disjunct, and occur only in a handful of sites under highly restricted environments in
this area. These occur on steep, north/northeast facing slopes at the far eastern edge of the escarpment,
on substrates dominated by fractured granite and having little soil development. Elevation range of the
limber forest type in the central eastern Sierra is 2590-3100 m. More dominant Sierran conifers abound
in the same elevation range as limber pine. At lower elevations, associates may include Jeffrey pine (P.
jeffreyi), western white pine (P. monticola), white fir (Abies concolor) and lodgepole pine (P. contorta); at
mid and upper elevations, red fir (A. magnifica), and mountain hemlock (Tsuga mertensiana) occur.
Whitebark pine (P. albicaulis) is the dominant Sierran subalpine species, and ranges from middle
elevations with limber pine upward to create the Sierran upper treeline at 3444 m.
The first tier of significant (over 3000 m) Great Basin ranges east of the Sierra Nevada in the focal region
includes, from north to south, the Sweetwater Mountains, the Bodie Mountains, and the Glass Mountain
Range. These ranges receive intermediate climates between Sierran and Great Basin, and similarly the
upper elevations (above pinyon/juniper woodland and sagebrush steppe) have forest vegetation that
derives from both bioregions. Limber pine in these ranges is much more abundant than in the Sierra
Nevada, occurs on many more slope aspects, and has a wider elevation range (2070 – 3400 m). Sierran
conifers occasionally occur; for instance, whitebark pine, lodgepole pine, and western white pine occur in
the Glass Mountains; white fir, lodgepole pine, and mountain hemlock occur in the Sweetwaters, but in
general these species are outliers in the western Great Basin, and highly restricted in extent.
The second tier of significant Great Basin ranges eastward includes, from north to south, the Wassuk
Range and the White/Inyo Mountains. Above the pinyon/juniper woodland in the Wassuk Range, limber
pine is the only conifer to form forest types; individuals of Jeffrey pine and whitebark pine have been
reported, but are rare and we have not documented whitebark pine from the range. Limber pine generally
grows in very sparse, open communities, limited to north and northeast aspects between 2665 and 3220
m. On Corey Peak, at the southern edge of the range, limber pine stands are unusually dense and occur
on north, northeast, northwest, east, southwest, and south facing aspects.
High-elevation conifers of the White/Inyo Mountains include Great Basin bristlecone pine and limber pine.
Both species co-occur at upper treeline, which extends to 3505 m. Bristlecone pine has a lower elevation
limit generally at 3040 m, although occasional outlier stands are lower, while limber pine commonly
extends down to 2665 m.
Contemporary Climate
The current climate of the focal region extends from California montane Mediterranean climate at the
Sierra crest on the western edge to interior, continental montane climate with monsoon influence at the
eastern side. The Mediterranean climate is characterized by precipitation that occurs primarily in winter
from Pacific frontal storms that move eastward depositing snow orographically in the western and high
Sierra. Snowfall is high near the Sierra crest and decreases rapidly eastward along the escarpment.
Gradients of precipitation and temperature are steep and the eastern slopes of the Sierra Nevada lie in a
rainshadow. Summer temperatures range from warm/moderate at high elevations to hot and dry in the
basins immediately east of the crest; atmospheric humidities are low throughout the region, although
higher at upper elevations. Moving eastward from the Sierran escarpment, precipitation is lowest
immediately east of the Sierra Nevada, with the low-lying basins (average elevation from 1200 – 2130m)
being the driest locally. Several upland regions extend as east-west ridges and serve to connect interior
ranges climatically with the Sierra Nevada. These include the Bodie Mountains and the Glass Mountain
Range, north and south, respectively, of Mono Lake, and allow the passage of cool, moist air from the
Sierra Nevada to the Sweetwater Mtns and Glass Mountain, respectively.
Further east, basin lowlands isolate the mountain ranges, and the White/Inyo and Wassuk ranges are
continental islands climatically. Owing to general circulation patterns in western North America as well as
their continental geography and location relative to the Gulf of California, the eastern mountains of the
region have climates more akin to the Rocky Mountains than to the Sierra Nevada. Winters are cold and
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relatively dry, with lower snowpacks than the Sierra; summers receive moisture from both local
convection storms and from gulf-origin monsoon storms, which reach their northern limit in this area.
The overall gradients of these regional climate patterns are mitigated by complex topography, so that
some localities receive more winter and summer storm activity while others of equal bulk and elevation
receive less.
Weather stations with relatively long periods of record in and adjacent to the region include: Bishop, CA,
Yosemite Valley, CA, Lee Vining, CA, Bridgeport, CA, Woodfords, CA, Kirkwood, CA, Carson/Reno, NV,
and Mina, NV
Paleoclimate
Paleoclimates are relatively well documented in the eastern Sierra/western Great Basin region due to the
diverse proxies available and favorable conditions for preservation of records, but significantly these are
at a coarse, region-wide scale. Long-lived climate-sensitive conifers contain multi-millennial tree-ring
records that preserve in dry, cold environments (e.g., Graumlich, 1993; Lloyd, 1997); semi-arid
environments support habitat for packrats, whose middens provide long historical records of desert
vegetation (Thompson, 1990; Grayson, 1993). Ancient pluvial as well as abundant montane lakes have
provided sediments for pollen, diatom, charcoal, and physical paleo-analysis (Anderson, 1990; Li et al.,
2000; Benson et al., 2002). The neoglacial advances of the high Sierra Nevada document mountain
climate over the last several thousand years (Birman, 1964; Clark and Gillespie, 1997), and hydrologic
analysis of river and lake levels (Stine, 1990, 1994) provide a historic baseline of drought, snowpack, and
water availability. Volcanism within and adjacent to the Long Valley Caldera system has been active in
the late Holocene, especially among of the vents of the Inyo and Mono Craters Chains (Miller, 1984,
1985). Not only have these contributed to landscape evolution directly, but regular ash eruptions have
unique chemical signatures that have enabled accurate regional stratigraphic controls on dating
The time depth relevant to limber pine records begins about 4000 years ago. This period is significant as
it marks the first general cooling trend after the peak heat and extensive dry of the middle Holocene
(6000-4000 years ago). The centuries beginning about 4000 years ago have been characterized as the
neoglacial or neopluvial period (Konrad and Clark, 1998, Stine, 1994, Tausch et al., 2004). Glaciers
advanced in the Sierra Nevada for the first time since the end of the Pleistocene, and the climate
generally cooled from a mid-Holocene interglacial warm peak and became much wetter (LaMarche,
1974). Treelines generally lowered, woodlands increased at mid- to low elevations, and desert shrub
vegetation decreased. This extensive period was \ followed by a severe extended drought, known as the
post-neoglacial drought (Tausch et al., 2004), which lasted from about 2500 – 1300 years ago. During
this time, diverse proxies indicate a significant drop in precipitation, with resulting decreases in woodland
and forest vegetation, increase in desert shrubs, and significant slope erosion, although temperatures
were likely cooler than the mid-Holocene.
The period from ca. 900 – 1350 CE is known as a time of significant global climate change -- the
Medieval Climatic Anomaly (MCA). Whereas many parts of the world experienced deviations up to 1°C
warmer during this time, the primary effect in the Sierra Nevada appears to have been decreased
available moisture for plants, although it is not clear what the interaction with temperature and snowpack
retention was. Stine documented two extensive droughts, at AD 900-1112 and AD 1200-1350 (Stine
1994), evident in the central and eastern Sierra Nevada. Climate models derived from tree-ring variability
in the southern Sierra Nevada indicate that temperatures were slightly warmer than average during this
time (Graumlich, 1993). Our work modeling climates based on the intersection of climate space for the
species found on Whitewing Mtn during the MCA indicates that, relative to present, the MCA period was
much warmer especially in winter minimum temperatures and slightly wetter, especially in winter (Westfall
and Millar 2004; Millar et al., in prep). Although ecosystem responses in the Sierra Nevada to MCA have
not been widely studied, those documented suggest changes in fire regime (Swetnam, 1993), treeline
elevation, and tree growth (Graumlich and Lloyd, 1996), and increases in abundance of firs (Abies) in
high-elevation communities (Anderson, 1990).
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The MCA ended abruptly in the late 1300s, as climates cooled globally during a five-century period (AD
1400-1900) known as the Little Ice Age (LIA). In the Sierra Nevada significant glacial advances (Clark
and Gillespie, 1997; Konrad and Clark, 1998) and reversals in some of the ecosystem reponses (Lloyd
and Graumlich 1997; Anderson 1990; Swetnam, 1993) characterized the LIA. Twentieth century warming
has been widely documented worldwide as well as in western North America and the Sierra Nevada
(Millar et al., 2004a). In general, the century reflects a rebound from the LIA, with increasing
temperatures, especially minimum temperatures, and increased precipitation. Notable also in the 20thcentury are increases in extreme climate events, including those related to the ENSO cycle as well as
multi-decadal modes such as the Pacific Decadal Oscillation (Millar et al., 2004a, 2004b).
4. Objectives
The overall objective of the study is to evaluate medium- to high-resolution spatial and temporal response
of limber pine populations to climate variability at centennial to decadal scales over the past 4000 years
and across a geographic gradient from the Sierra Nevada to the western Great Basin.
Questions:
1. What are the age structures recorded in live and reconstructed (from deadwood) limber pine forests at
nested temporal (centennial and decadal) and spatial (mountain range, sub-range, region within range,
and local site) scales across the study region?
2. If demographic patterns are revealed, how do they relate to centennial- to decadal-scale climates for
the local regions as recorded from other proxies?
3. If demographic patterns are revealed, how do they compare among mountain ranges, watersheds, and
aspect/elevations?
4. At the high-resolution spatial scale, what is the relative role of change in treeline elevation in response
to climate variability versus change in stand aspect and structure?
5. What climatic reconstructions can be derived from resulting limber pine tree-ring chronologies?
6. What implications do these results have for present forest dynamics in regard to changing present and
future climates and in what specific ways would this influence management and policy decisions for midelevation forests?
5. Methods
Study Sites
The optimal design is spatially replicated and geographically hierarchic (table 1). The primary design is a
2 x 3 rectangular grid that includes three tiers of mountain ranges, from west to east with two replicate
areas at the north and south of each range (designated as 6 sub-ranges). Two regions (watersheds,
distinct summits or ridges) would be selected within each sub-range. Within each region 1-many sites
would be chosen. Optimal sites are watersheds that include limber pine distributed over a range of
aspects and elevations. Within each site, a minimum of 50 limber pine live and/or deadwood would be
sampled, optimally selecting stems distributed over elevation and aspect. Thus, a minimum of 600
samples would be collected, and optimally a minimum of 1200 samples. Table 1 lists the structure of the
study design, and includes initial site selections.
In addition to the main grid of sites, two satellite regions are added to the main study: the Toiyabe Mtns
and Owens Gorge (fig. 1 and table 1). We propose sites in the Toiyabe Mtns as a joint project with Dr.
Robin Tausch, Research Paleoecologist, USDA Rocky Mtn Research Station, Reno, NV. Tausch has
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worked extensively on paleoecology of mid-elevation woodlands in the Toiyabe Mtns, and our teams
have mutual interest in addition of limber pine studies in this area. Not only are the opportunities for
collaboration valuable, but the Toiyabe Mtns afford a useful outlier site to our main study, as climate and
floristics in the central Great Basin as present is quite different from our focal region.
The Owens Gorge sites are proposed as outlier low-elevation contrasts. Limber pine populations growing
in Owens Gorge, near Bishop, CA, are unusual for their context. They occur below the lower treeline of
pinyon pine in the area, and far below the dominant limber pine treeline regionally. Demographics of the
site also appear very different from common limber pine populations, as there is little deadwood in the
stand and the trees appear young and fast-growing; an undiagnosed insect or disease is afflicting the
stand at present. Dissecting the history and climate relations in this region should yield important
information on factors affecting limber pine’s growth that will assist interpretations in the main study.
Field Measurements
At each site, limber pine trees will be selected in a distribution to achieve for two purposes: to develop
conventional tree-ring chronologies (i.e., to enable cross-dating and for subsequent climatic
reconstruction) and to adequately sample demographic structure. For the former, a range of live and
dead trees occurring on severe sites under stress conditions is optimal. To maximize chances of
obtaining oldest wood for chronology building, deadwood samples will be sought in locations likely for
long preservation, i.e., rocky outcrops, ridgetops, other severe sites. For demographic sampling, live
trees and deadwood will be sampled as evenly distributed as possible across the range of elevations,
aspects, slopes, soil substrates, within limitations of available trees and wood and dependent on local
conditions at the site.
Minimally each tree will be sampled once for cross-dating and age determination. Either an increment
core will be taken or wood section sawn from the main stem of live trees or deadwood. In cases where a
single increment core obviously does not sample the full age range of a tree, additional cores will be
taken. Occasionally branch or root samples will be taken if the stem is not adequately preserved and the
sample is not better represented by other stems. Sampled stems will be labeled in the field and
photographed. Using a Geographic Position System instrument (Garmin Venture model), latitude,
longitude, and elevation for each sample will be recorded. Elevation will be cross-checked with an
atmospheric-pressure-based altimeter. Slope aspect in azimuth degrees will be recorded, and length and
width of each sample measured.
Lab and Statistical Analyses
Increment cores and wood sections will be processed following standard dendrochronological methods
(Stokes and Smiley, 1968) with each tree-ring series measured to the nearest 0.002 mm in the Albany
tree-ring laboratory. Site-specific tree-ring chronologies will be built by cross-dating continuous samples
collected at each site, and by reference to previously published chronologies extracted from the
International Tree Ring Data Base. In our previous studies, we have found limber pine to cross-date well
with Great Basin bristlecone pine, for which several reference chronologies exist with continuous and
greater temporal depth than we expect from limber pine. Resulting chronologies developed from the best
representative samples at each site will allow precision in cross-dating samples collected for demographic
analysis. We will attempt to establish exact calendar dating of all live and deadwood stem cross-sections
using a combination of dendrochronological techniques, including correlation and skeleton plot
techniques (Holmes et al., 1983, 1986; Cook and Kairukstis, 1990).
Once dates of samples have been established and verified chronologies built we will be able to do the
following analyses:
1. Mountain Range (3 ranges) and Sub-Range (6 sub-ranges) Scale
-- Climate reconstructions, correlating ring-widths against long records from instrumental data collected at
weather-recording stations and accessible from the Western Regional Climate Center
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-- Climate reconstructions, as above, but modeling current climate from PRISM (Daly et al., 1994) rather
than from instrumental data directly.
-- Climate modality and cyclicity: wavelet and spectral analysis (SAS Institute, 2002, 2004), moving
window Webster analysis for low-frequency variability (Legendre and Legendre, 1998), 100-year filter for
high-frequency variability (Wolfram, 2004)
-- For comparison of growth and climate reconstructions among mountain range and sub-range sites:
correlation analysis of individual chronologies (SAS Institute 2004, 2005), with focus on periods of
persistent synchrony and asynchrony among sites, and correlations of sub-range to latitude/longitude
influenced by synoptic climatology patterns.
2. Regions within Sub-Range Scale (2 regions within each range)
-- Correlation analysis of individual chronologies (SAS Institute 2004, 2005), with focus on periods of
persistent synchrony and asynchrony among sites, and on average ring-width and time depths.
3. Sites within Regions (1-7 sites per region)
-- Demographic analyses, including correlation/regression analyses of ring-series dates against latitude,
longitude, slope, aspect, elevation, and as appropriate, substrate and associated floristics (SAS Institute
2004, 2005).
-- Correlation of occurrences of fire sign (charcoal) with latitude/longitude, elevation, slope, and aspect.
-- Estimation of upper and lower treeline over time at each site, based on moving window Webster
method (Legendre and Legendre, 1998)
6. Application of Research Results
We expect outcomes from the proposed work both to basic science and landscape-scale management
and conservation. We anticipate that evaluating the role of climate in influencing forest structure and
composition will improve understanding of forest dynamics and thus prescriptions for management
generally. The role of climate as an ecological architect has been especially difficult to comprehend and
assess at landscape- to stand-level scales. Management efforts have focused mostly on using results
from downscaled climate-change models as inputs to vegetation models to assess impacts of future
climate on forest growth. While such an approach yields clear quantitative answers, the results depend
on equilibrium conditions and linear changes in both climate and vegetation response. We anticipate that
our limber pine research will show these to be grossly inadequate assumptions, and that instead,
management projections and policy for the future must take into account the nested modes of variability
likely in the future, and the diverse non-equilibrium responses of vegetation. We will attempt to translate
the results from our research into relevant lessons for forest management.
We anticipate a minimum of three technical publications from this work, to be submitted to Quaternary
Research Paleoecology, Paleoclimatology, and Paleobotany, and/or Ecology. Additional opportunities to
present the work orally and in posters at appropriate scientific meetings will be sought. Implications of
forest dynamics and relative roles of climate and vegetation response will be communicated into
management and conservation guidelines through journals and other outlets more appropriate to a
management audience, such as Ecological Applications.
7. Safety and Health
Standard procedures determined by the SNRC Safety Committee will be followed (see SNRC intranet).
Field and Office Job Hazard Analyses and Emergency Evacuation Procedures on file for Millar research
team pertain to and adequately cover safety procedures for this project.
8. Environmental analysis considerations (FSM 1950).
None applicable.
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Local Contacts
Hawthorne Amy Base: John Peterson, Resource & Facilities Manager
USFS Inyo National Forest: Larry Ford, Mono Basin Scenic Area Resource Manager, and Molly Brown,
Mono Lake District Ranger
USFS Toiyabe National Forest, Kathleen Lucich, Bridgeport District Ranger
9. Personnel Assignment, Time of Completion, and Cost
Millar, Principal Investigator. Oversight and supervision for all aspects of the study, including project
design, justification, grant applications, field work, data analysis, communication, quality control, field and
office safety, and publication/dissemination.
Westfall: Principal Co-Investigator. Primary input on field and statistical design, analysis, and statistical
interpretation. Assists and reviews study plan, participates in and advises on field techniques and lab
analyses, provides input and review on manuscripts.
Delany: Laboratory Analyses (ring measurements, data input, basic statistic analyses), and assists in
development of graphics for publications, posters, oral presentations.
King: Field Assistance (mapping, measurements) and assistance/input to ring measurement of difficult
sections, interpretation of ring measurement results for dendroclimatic analyses.
Completion Dates:
Fieldwork: October 2005
Lab Analyses: March 2006
Statistical Analysis: June 2006
Manuscript Preparation & Review: Winter 2007
Remaining cost: Salary for Millar, Westfall & Delany; King donates time to this project, and occasionally
works on small purchase orders for specific contributions.
10. References and Additional Bibliography
Anderson, R.S. 1990. Holocene forest development and paleoclimates within the central Sierra Nevada,
California. J. Ecol. 78: 470-489.
Benson, L. Kashgarian, M., Rye, R., Lund, S., Paillet, F., Smoot, J., Kester, C., Mensing, S., Meko, D.,
Lindstrom, S., 2002. Holocene multidecadal and multicentennial droughts affecting northern California
and Nevada. Quaternary Science Reviews 21, 659-682.
Cheng, S. (ed.) 2004. USDA Forest Service Research Natural Areas in California. USDA Forest Service,
General Technical Report, PSW-GTR-188. Albany, CA: Pacific Southwest Research Station, Forest
Service, U.S. Department of Agriculture; 338p
Clark, D.H. and A.R. Gillespie. 1997. Timing and significance of late-glacial and Holocene cirque
glaciation in the Sierra Nevada, California. Quaternary Research 19:117-129.
Clark, J.S. and Hussey, T.C. 1996. Estimating the mass flux of charcoal from sedimentary records:
Effects of particle size, morphology, and orientation. Holocene 6(2): 129-144.
Cook, E.R., Holmes, R.L., 1992. Program CRONOL, in: H.D. Grissino-Mayer, R.L. Holmes, H.C. Fritts
(Eds.), International Tree-Ring Data Bank Program Library, user’s manual, Laboratory of Tree-Ring
Research, Tucson (AZ).
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Daly, C., R.P. Neilson, and D.L. Phillips. 1994. A statistical-topographic model for mapping climatological
precipitation over mountainous terrain. Journal of Applied Meteorology 33, 140-158.
Graumlich, L.J. 1993. A 1,000 year record of temperature and precipitation in the Sierra Nevada.
Quaternary Research 39: 249-255.
Graumlich, L.J. and A. H. Lloyd. 1996. Dendroclimatic, ecological, and geomorphological evidence for
long-term climatic change in the Sierra Nevada, USA. Radiocarbon 51-59.
Grayson, D.K. 1993. The Desert’s Past. A Natural Prehistory of the Great Basin. Smithsonian Institution
Press. Washington. 356 pgs.
Holmes, R.L. 1983. Computer-assisted quality control in tree-ring dating and measurement. Tree-Ring
Bulletin 43: 69-78.
Holmes, R.L., Adams, R.K., Fritts, H.C., 1986. Tree-ring chronologies of western North America:
California, Eastern Oregon, and Northern Great Basin with procedures used in the chronology
development work including user’s manuals for computer programs COFECHA and ARSTAN. Laboratory
of Tree-Ring Research, University of Arizona, Chronology Series VI.
Konrad, S., Clark, D.H., 1998. Evidence for an early Neoglacial advance from rock glaciers and lake
sediments in the Sierra Nevada, California, U.S.A. Arctic and Alpine Research 30, 272-284.
LaMarche, V., 1973. Holocene climatic variations inferred from tree line fluctuations in the White
Mountains, California. Quaternary Research 3, 632-660.
Legendre, P., and Legendre, L., 1998: Numerical Ecology..Second English Edition. Elsevier, New York.
Li, H.C., Bischoff, J.L., Ku, T.L., Lund, S.P., Stott, L.D., 2000. Climate variability in East-Central California
during the past 1000 years reflected by high-resolution geochemical and isotopic records from Owens
Lake sediments. Quaternary Research 54, 189-197.
Lloyd, A.H., 1997. Response of tree-line populations of foxtail pine (Pinus balfouriana) to climate variation
over the last 1000 years. Canadian Journal of Forest Research 29, 936-942.
Millar, C.I. 2004. Climate change as an ecosystem architect: Implications to rare plant ecology,
conservation, and management. In, Brooks, M. (ed). Proceedings of the Conference on Rare Plants,
Ecology, and Conservation, 9-13 Feb 2002, Arcata, California. California Native Plant Society. Pgs 139157.
Millar, C.I. and L.B. Brubaker. In press. Climate change and paleoecology: New contexts for restoration
ecology. In M. Palmer and D. Falk (eds) Restoration Science. Island Press.
Millar, C.I., R.D. Westfall, D.L. Delany, J.C. King. In prep. WW mss
Millar, C.I., R.D. Westfall, D.L. Delany, J.C. King, and L.C. Graumlich. 2004a. Response of subalpine
conifers in the Sierra Nevada, California, U.S.A., to 20th-century warming and decadal climate variability.
Arctic, Antarctic, and Alpine Research 36 (2): 181-200.
Millar, C.I., R.D. Westfall, D.L. Delany, J.C. King and Alden, H.A.. 2004b. Late Holocene dynamics of the
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Table 1. A. Main Study. Locations and hierarchic structure of optimal study sites, with initial selections,
for limber pine across a gradient of three mountain ranges from the Sierra Nevada to the western Great
Basin. Optimal structure is 3 tiers of mountain ranges, from west to east and 2 replicates north and south
(= sub-ranges); 2 regions within each sub-range; and 1-2 sites within regions. Optimal sites within
regions are those that include a range of elevations and aspects.
A. Main Study
Mountain Range
and Sub-Range
Region within
Sub-Range
Site within
Region
1st Tier (West)
Sierra Nevada N
Kavanaugh
Lundy
1
1
Parker
Glass Cr
1
1
Sweetwater Cyn
Silverado Cyn
2
1
Glass Mtn
Kelty Ridge
1
2
Mt Grant
Corey Peak
7
2
Trail Canyon
Middle Fk Canyon
3
1
N. Toiyabe Peak
N San Juan
1
1
N Slope
4
Sierra Nevada S
2nd Tier (Intermediate)
Sweetwater Mtns
Glass Mtns
3rd Tier (East)
Wassuk Range
White Mtns
B. Satellite Regions
Toiyabe Mtns
Owens Gorge
13
Figure 1. Map of focal research region and individual study sites for
climate research on limber pine in the eastern Sierra Nevada and western
Great Basin. See Table 1 for site names.