Ecological Setting of the Wind River
Old-growth Forest
David C. Shaw,1,2* Jerry F. Franklin,1,2 Ken Bible,1,2 Jeffrey Klopatek,3
Elizabeth Freeman, Sarah Greene,4 and Geoffrey G. Parker5
1Wind River Canopy Crane Research Facility, University of Washington, Carson,3 Washington 98610, USA; 2College of Forest
Resources, University of Washington, Seattle, Washington 98195-2100, USA; 3Department of Plant Biology, Arizona State University,
Tempe, Arizona 85287-1601, USA; 4Pacific Northwest Research Station, Forestry Sciences Laboratory, US Forest Service, Corvallis,
Oregon 97331, USA; 5Smithsonian Environmental Research Center, Edgewater, Maryland 21037-0028, USA
ABSTRACT
The Wind River old-growth forest, in the southern
Cascade Range of Washington State, is a cool (av-
erage annual temperature, 8.7
C), moist (average
annual precipitation, 2223 mm), 500-year-old
Douglas-fir4 ?western hemlock forest of moderate to
low productivity at 371-m elevation on a less than
10% slope. There is a seasonal snowpack (Novem-
ber?March), and rain-on-snow and freezing-rain
events are common in winter. Local geology is
characterized by volcanic rocks and deposits of
Micocene/Oligocene Micocene-Oligocene5 (mixed)
Micocene and Quaternary age, as well as intrusive
rocks of Miocene age. Soils are medial, mesic, Entic
Vitrands6 that are deep (2?3 m), well drained, loams
and silt loams, generally stone free, and derived
from volcanic tephra. The vegetation is transitional,
between the Western Hemlock Zone and the Pacific
Silver Fir Zone, and the understory is dominated by
vine maple, salal, and Oregon grape. Stand struc-
tural parameters have been measured on a 4-ha
plot. There are eight species of conifers, with a stand
density of 427 trees ha)1 and basal area of 82.9 m2
ha)1. Dominant conifers include Douglas-fir (35
trees ha)1), western hemlock (224 trees ha)1), Pa-
cific yew (86 trees ha)1), western red cedar7 (30 trees
ha)1), and Pacific silver fir (47 trees ha)1). The av-
erage height of Douglas-fir is 52.0 m (tallest tree,
64.6 m), whereas western hemlock averages 19.0 m
(tallest tree, 55.7 m). The regional disturbance re-
gime is dominated by high-severity to moderate-
severity fire, from which this forest is thought to
have originated. There is no evidence that fire has
occurred in the forest after establishment. Primary
agents of stand disturbance, which act at the indi-
vidual to small groups of trees scale, are wind, snow
loads, and drought, in combination and interacting
with root-rot and butt-rot fungi, heart-rot fungi,
dwarf mistletoe, and bark beetles. The forest com-
position is slowly shifting from dominance by
Douglas-fir, a shade-intolerant species, to western
hemlock, western red cedar, Pacific yew, and Pacific
silver fir, all shade-tolerant species. The Wind River
old-growth forest fits the regional definition of
Douglas-fir ??old growth?? on western hemlock sites.
Key words: old growth; temperate coniferous
forest; ecological characterization; soils; geology;
climate; vegetation; disturbance.
INTRODUCTION
The Wind River old-growth forest is an approxi-
mately 500-year-old Douglas-fir (Pseudotsuga men-
ziesii)?western hemlock (Tsuga heterophylla) forest
in the southern Cascade Range of Washington
State. The Wind River Canopy Crane Research
Facility (WRCCRF) was established in this forest to
provide access to the canopy (Figure 1). The crane
is a 75-m-tall construction tower crane with an 85-
m jib that provides three-dimensional access to 1.7
? 106 m3 of the canopy over a 2.3-ha ??canopy
crane circle?? via a gondola (personnel basket)
Received 15 February 2002;1 accepted 31 October 2002; published online
12 May 2004.
*Corresponding author; e-mail: dshaw@u.washington.edu
Ecosystems (2004) 7: 427?439
DOI: 10.1007/s10021-004-0135-6
427
suspended from the jib. The College of Forest Re-
sources, University of Washington, USDA Pacific
Northwest Research Station (Portland, Oregon,
USA) and USDA Gifford Pinchot National Forest
(Vancouver, Washington, USA) jointly established
the facility in 1995.
In 1997, the Wind River old-growth forest and
the WRCCRF were selected as a primary study site
for the investigation of forest carbon dynamics by
the Western Regional Center (WESTGEC) of the
National Institute for Global Environmental
Change. Eddy-flux instrumentation was estab-
lished on the tower of the crane in 1998 at which
time the WRCCRF became an AmeriFlux site. This
report provides an overview of the physical and
ecological setting of the Wind River old-growth
forest in order to better understand, interpret, and
extrapolate the results of the detailed process-based
research reported elsewhere in this special feature8 .
DESCRIPTION OF THE FACILITY
The quantified forest descriptions provided in
this report are based on a 4-ha stem mapped plot
that surrounds the canopy crane (WRCCRF 4-ha
plot). The site is located in the Wind River valley of
the southern Washington Cascade Range, approx-
imately 75 km east of Portland, Oregon, near Car-
son, Washington, and located at 371-m elevation,
at latitude N 45
49?13.76?? and longitude W
121
57?06.88??. The valley emerges into a major
geographic feature, the Columbia River Gorge,
about 15 km south of the crane site (Figure 2).
Administratively, the Wind River old-growth forest
and the canopy crane are located within the T. T.
Munger Research Natural Area, and the Trout
Creek Division of the Gifford Pinchot National
Forest?s Wind River Experimental Forest. The Wind
River Experimental Forest (WREF) is a 4208-ha
area formally dedicated to scientific and educa-
tional purposes by the USDA Forest Service in 1932
and has been an active forest research site since
1908. The T. T. Munger Research Natural Area
(RNA) is a 478-ha old-growth forest preserve
(about 500 years old) within the WREF and is
dominated by Douglas-fir and western hemlock.
The RNA was selected for protection in 1926 and
expanded in size in 1934, as an area representative
of the widespread old-growth Douglas-fir forests
found west of the crest of the Cascade Range in
Washington and Oregon (Franklin 1972). USDA
Forest Service Pacific Northwest Research Station
and Gifford Pinchot National Forest are jointly re-
sponsible for administration of the Experimental
Forest and Research Natural Area.
NATURAL AND HUMAN GEOGRAPHY
The defining geographic feature of the Wind
River old-growth forest area is the north?south-
oriented Cascade Range and the Columbia River
Gorge, 15 km south, which provides a water-grade
east?west pathway through the range. The Cascade
Range extends from northern California to south-
ern British Columbia, with a width of 75?100 km
in the central region. Major divides are typically
1000- to 2000-m elevation, with volcanic cones
occasionally reaching 3500-m elevation. The Cas-
cade Range is a major barrier to the primarily
easterly flow of marine air masses from the Pacific
Ocean. Consequently, clouds are abundant and
rainfall increases rapidly on the western slopes of
Figure 1. Vertical profile through the old-growth forest at the Wind River Canopy Crane Research Facility (WRCCRF).
The drawing includes all trees larger than 5-cm diameter at breast height within a swath 180 m long and 10 m wide
through the center of the WRCCRF 4-ha plot. The crane is 75 m tall with an 85-m-long jib, accessing 2.3 ha of ground area
under the swing of the jib. The stand is dominated by Douglas-fir in terms of height, wood volume, and basal area, but
western hemlock in terms of number of trees.
428 D. C. Shaw and others
the range due to orographic lift; the rapid increase
with elevation on the western slopes is mirrored by
a strong rain-shadow effect on the eastern slopes.
The Columbia River Gorge provides a natural
pathway for the movement of air masses and or-
ganisms between the moist, moderate maritime
environment west of the Cascade Range and the
arid, more extreme continental environment east
of the range. This circumstance results in some
distinctive climatic events (Lawrence 1939). For
example, the dominant westerly, year-around
winds that are associated with mild marine air
masses are occasionally interspersed by periods
with strong easterly winds that bring dry cold (in
winter) or hot (in summer) continental air from
the interior. In addition, periodic collisions be-
tween Pacific storms and cold continental air result
in severe ice, rain-on-snow, and snow storms.
The Wind River old-growth forest is located
within the Wind River valley 9on gentle topography
(less than 10% slope) between two major topo-
graphic features?Trout Creek Hill and Bunker Hill
(Figure 2). The crane and WRCCRF 4-ha plot are
actually located on the southeastern foot of Trout
Creek Hill (Figure 2), a shield volcano with a
summit of 792-m elevation. Bunker Hill (636-m
elevation) is a prominent steep-sided feature about
1 km to the southeast that affects the crane site by
delaying direct sunlight for an hour or so in the
morning and by providing some protection from
east winds. The walls of Wind River valley itself rise
approximately 1500 m above the valley floor, with
the nearest located approximately 2 km to the
south. The crane facility is located near the subtle
divide between the Wind River and Trout Creek, a
major tributary of the Wind River. The location of
the forest near the divide between two watersheds
limits the extent of soil water available to the site
and contributes to dry soils during summer. Cold
air drainage is primarily from the north, as higher-
elevation hills contribute cooling. The site is pro-
tected from severe windstorms 10, which come pri-
marily from the west due to the north?south
orientation of the valley bottom.
There have been 11significant human impacts on
environmental conditions of the Wind River old-
growth forest. The most important was the clearing
of the forest and establishment of tree seedling
nursery fields along the southern boundary of the
RNA in the mid-1980s. This resulted in creation of a
sharp, high-contrast edge 50?60 m high. The
microclimatological consequences of this edge have
been documented by Chen and colleagues (1992,
1993a, 1993b). Edge influences were found to ex-
tend into the forest for 200 m. The crane is located
approximately 500 m from this edge and, therefore,
is not strongly impacted. However, the proximity to
the fields does appear to affect some vertebrate spe-
cies, such as elk, which bed in the old growth and
feed in the fields. The influence of this edge on me-
teorologic conditions and the eddy-flux stations is
discussed elsewhere (Paw U and others 2004). The
80-year-old Douglas-fir forest 500 m to the north of
the crane site was logged around 1920 and was
thinned in the early 1990s, but still provides a nearly
intact buffer for the natural area and crane facility.
CLIMATE
The Wind River old-growth forest climate sum-
mary is based on the 1978?98 National Oceano-
graphic and Atmospheric Administration (NOAA)
National Climate Data Center climate data
(www.ncdc.noaa.gov) collected at the Carson Na-
tional Fish Hatchery (US Fish and Wildlife Service)
located 5 km north of the WRCCRF. Annual pre-
cipitation has historically averaged 2223 mm y)1
(SD = 5.7), with only about 5% falling during
Figure 2. Area relief map of the Wind River Valley, in-
cluding Trout Creek Hill (792-m elevation), Bunker Hill
(636-m elevation), Wind River Canopy Crane Research
Facility (WRCCRF, 371-m elevation), and the Columbia
River (Lake Bonneville) (31-m elevation). The Wind
River valley bottom bedrock is composed of lava flows
from Trout Creek Hill, a Quaternary volcano approxi-
mately 340,000 years old. Bunker Hill is composed of
Miocene gabbro intrusive rock underlain by volcano-
clastic deposits of Miocene/Oligocene age.
Wind River 2Old-growth Forest 429
June, July, and August (Figure 3). This summer
drought is an important characteristic of Pacific
Northwest climate because it profoundly reduces
summertime tree growth (Franklin and Waring
1980). Much of the winter precipitation falls as
snow, and snowpack (depth) is over 100 mm dur-
ing December, January, February, and March
(Figure 3). Average annual air temperature was
8.7
C (SD = 6.5). Average monthly air temperature
in January was 0.1
C (SD = 2.3), while average
monthly air temperature in July was 17.7
C
(SD = 1.7).
The Wind River old-growth forest is in the tran-
sient snow zone of the Cascade Mountains, where
rain-on-snow, freezing-rain, and graupel events are
common. The importance of being in this transi-
tional zone of winter climate cannot be overstated.
The heavy buildup of wet snow on tree branches
has a major influence on branch and bole breakage.
For example, over 50% of the western hemlock
trees in the crane site have evidence of leader and
bole breakage. The deep, complex structured can-
opy is expected to play a major role in modifying the
amount and timing of snowmelt runoff in this zone;
old-growth forests reduce the peak flows associated
with rain-on-snow events compared with cleared or
very young forests (Harr 1986).
GEOLOGY AND LOCAL GEOGRAPHY
The Wind River valley is characterized by vol-
canic rocks and deposits of Miocene/Oligocene and
Quaternary age as well as intrusive rocks of Mio-
cene age (Walsh and others 1987). Further geo-
logical information on the area can be found in
Wise (1961, 1970), Hammond (1980), and Ham-
mond and Korosec (1983). Although glacial de-
posits dated at more than 38,000 years before
present have been identified in the region, details
of glaciation are not well understood. Trout Creek
Hill (Figure 2), the geologic feature on which the
old-growth forest developed, is a Quaternary vol-
cano approximately 340,000 years old. A cinder
cone and lava tunnels are located at the summit.
Dark-gray olivine basalt flowed from Trout Creek
Hill (and possibly other vents) down the Wind
River Valley in a series of intracanyon lava flows
that reached the Columbia River (Walsh and others
1987; Woodfin and others 1987). These flows make
up the bedrock that underlies the research site, as
well as much of the valley floor. Bunker Hill is a
prominent landform composed of Miocene gabbro
intrusive rock underlain by volcanoclastic deposits
of Miocene/Oligocene age.
The old-growth forest and crane site are located
on an alluvial fan from Trout Creek Hill (Fig-
ure 2). The fan is slightly dissected by small drai-
nages. Total relief across the crane circle is about
10 m, with the lowest areas on the north and east
and the highest on the west and south. An
ephemeral stream runs through the crane circle
about 30 m north of the crane. The creek is most
deeply incised (about 2 m) on the west. After
leaving the crane circle, the stream flows into a
forested wetland to the northeast of the crane. The
wetland is located at the base of Bunker Hill and
drains 2 km to the north (as Cold Creek) into
Wind River.
SOILS
The soils are classified as medial, mesic, Entic
Vitrands12 (Stabler Series, www.statlab.iastate.edu/
cgi-bin/osd/osdname.cgi). These soils are deep, well
drained, generally stone free, and medium tex-
tured. The parent material is air-deposited, mixed
volcanic tephra, of relatively recent origin (be-
tween 3500 and 12,000 years before present). A
typical soil profile has a mull forest floor (average
depth, 6 cm; SD = 2.1; n = 74) above the mineral
soil. The surface mineral layer is generally 45-cm-
thick, dark yellowish brown shotty loam with ap-
proximately 5%?8% clay content. Volcanic mate-
rial, ranging from 2 to 10 mm in size, accounts for
40%?50% by volume of the upper soil. The subsoil
is commonly 45- to 110-cm-deep, brown or yel-
lowish brown silt to clay loam. The soil from 110 to
Figure 3. Climate regime from 1978 to 1998 for average
monthly air temperature and average monthly precipi-
tation, and total precipitation and snowpack (depth).
Data were collected at the Carson National Fish Hatchery
(www.ncdc.noaa.gov) located approximately 5 km north
of the Wind River old-growth forest.
430 D. C. Shaw and others
130 cm in depth is yellowish brown to strong
brown massive clay loam. Some soil profiles exhibit
up to 15% pumice gravels and coarse sand in lower
soil layers. Abrupt depositional discontinuities are
encountered at the lower depths that contain large
amounts of gravels and cobbles, some of which are
waterworn. Plant roots are concentrated above
50 cm in soil profiles; however, roots as deep as
2.05 m have been observed in younger forests
growing on nearly identical soils (T. Hinckley per-
sonal communication). Many coarse roots of
Douglas-fir extend to depths greater than 1.0 m.
Tip-up mounds of windthrown western hemlock
trees typically have a classic flat root plate indica-
tive of shallow rooting.
The mineral soils have a low bulk density,
characteristic of Andisols with high organic matter
content (Brady and Weil 1998; Cromack and others
1999). Bulk density of the top 20 cm of soil ranged
from 0.70 to 0.92 g cm)3, increasing with depth.
Soil pH (water to soil, 2:1) ranged from 4.9 to 5.7,
values typical for Douglas-fir forests in this region
[for example, see Heilman (1981)]. Organic matter
contents (calculated as percent total C/0.58) ranged
from 4.6% to nearly 10%. Values for carbon (C)
and nitrogen (N) concentrations of the forest floor
and mineral soil (Table 1) are comparable to most
other similar forests in the region [for example, see
Youngberg (1966) and Means and others (1992)]
and are generally in the midrange for forest soils in
the region. Based on 499 soil pedons in western
Oregon, soil C for the upper 20 cm averaged 6.5 kg
m)2 (range, 0.9?24) (Homan and others 1995),
whereas the canopy crane site has 9.7 kg m)2 in
the upper 20 cm of soil. In another study of soils
under four Douglas-fir?western hemlock forests
(Radwan 1992), N concentrations in the forest floor
(Oi, Oe, and Oa13 ) averaged from 0.78% to 1.06%,
while N content of the upper 20 cm of soil ranged
from 0.09% to 0.46%. The forest floor values of
the WRCCRF 4-ha plot fall within these ranges
(Table 1). The soil concentrations are reflective of
the less than 2.00-mm portion of soil, although the
larger tephra (mentioned previously) contained
both C and N concentrations approaching those of
the less than 2.00-mm fractions of soil (Cromack
and others 1999). These large particles are known
to have a high retention of phosphorus and
absorptivity of organic C and N (Brady and Weil
1996).
The pattern of C and N across the research site
reflects the microsite soil heterogeneity of the old-
growth forest system, a result of windthrow
mounds and large, buried, coarse woody debris.
Coarse woody debris constitutes over 93 Mg C ha)1
(Harmon and others 2004). The northeast quadrant
of the WRCCRF 4-ha plot is seasonally inundated
by water, and this results in a greater storage of C
and N, presumably due to the seasonal anaerobic
conditions and reflected in seasonal patterns of CO2
efflux (J. Klopatek unpublished14 ). Since December
1998, the water table has been monitored weekly
with four peziometers running across the slope at
the site. In the downhill (northeast quadrant of the
WRCCRF 4-ha plot) well, from December through
March the water table is highest and varies from 50
to 30 cm below the surface. In contrast, from
September through October the water table is
lowest and varies from 200 and 235 cm below the
surface. In the uphill (southwestern quadrant of
the WRCCRF 4-ha plot) well, from December
through March the water table varies from 206 to
188 cm below the surface, whereas from May
through October the water table drops below the
well at 234 cm.
UNDERSTORY VEGETATION
The WRCCRF plot is mapped on the border of the
Western Hemlock/Salal (Gautheria shallon)?West-
ern Hemlock/Oregon grape (Berberis nervosa)/Salal15
plant association complex and Pacific Silver Fir
Table 1. Concentrations of Carbon (g C kg)1 Dry Soil) and Nitrogen (g N kg)1 Dry Soil) and C:N Ratios in
the Forest Floor and Soils from the Wind River Canopy Crane Research Facility 4-ha Plot
Component Carbon g kg)1 Nitrogen g kg)1 C/N
Forest floor
Oi 452.9 (10.2) 8.0 (0.4) 59.7 (3.5)
Oe and Oa 436.6 (11.8) 9.4 (0.4) 48.56 (2.6)
Soil
0?10 cm 52.6 (3.7) 1.9 (0.1) 28.0 (1.2)
10?20 cm 33.8 (2.1) 1.4 (0.1) 24.6 (1.1)
Values are the mean (n = 24) and standard errors (in parentheses). All material was analyzed for C and N on a Perkin-Elmer 2400 CHN33 analyzer (Perkin-Elmer, Norwalk,
CT, USA). Oi, Oe, Oa34 , 0?10 cm, and 10?20 cm, respectively.
Wind River 2Old-growth Forest 431
(Abies amabilis)/Salal plant association (Meyers and
Fredricks 1993). These plant associations, charac-
terized by a dominance of Salal and Oregon grape,
are very widespread in the southern Cascades of
Washington, especially on ridges and upper slopes,
and are characterized by moderate to low produc-
tivity, with dry well-drained soils (Franklin and
Dyrness 1973; Topik and others 1986). However,
the abundance of winter snowpack, and a well-
developed component of Pacific silver fir indicate
that this site exhibits environmental conditions
approaching that of the Pacific Silver Fir Zone
(Brockway and others 1983).
Understory vegetation has an estimated leaf-area
index (LAI) of 1.69 (Thomas and Winner 2000),
which is 20% of the estimated total stand LAI of
8.61. We quantified understory plant cover and
frequency on sixty-four 25- ? 25-m subplots in the
WRCCRF 4-ha study area (Table 2). To date, we
have identified 68 vascular plant species (Tables 2
and 3), whereas Kemp and Schuller (1982) iden-
tified 178 species of vascular plants in the T. T.
Munger RNA, which includes wetlands to the east
of the crane site. Among the vascular plants, 33
plant families are represented, 18 by only one
species, and 7 by two species. Dominant plant
families include Ericaceae (8 sp.), Liliaceae (7 sp.),
Pinaceace (6 sp.), Polypodiaceae (5 sp.) and Rose-
aceae (5 sp.). Prominence values (PVs), determined
by multiplying the average percent cover by the
average percent frequency, indicate the dominant
understory species are vine maple (Acer circinatum,
PV 25.5), salal (PV 16.0), Oregon grape (PV 14.2),
vanilla leaf (Achlys triphylla, PV 4.4), red huckle-
berry (Vaccinium parvifolium, PV 4.2), inside-out
flower (Vancouvaria hexandra, PV 1.4), bracken16 fern
(Pteridium aquilinum, PV 1.4), twinflower (Linnaea
borealis, PV 1.1), and bear grass17 (Xerophyllum tenax,
PV 1.0) (Table 3). No other plant species rated a PV
1 or above.
A total of seven plant associations were keyed
from the 64 subplots (Figure 4) using a report by18
Topik and colleagues (1986). These include, from
wet-site to dry-site indicators, Western Hemlock/
Lady Fern (Athyrium filix-femina) (two subplots),
Western Hemlock/Foamflower (Tiarella trifoliata)
(four subplots), Western Hemlock/Alaska Huckle-
berry (Vaccinium alaskaense)-salal (two subplots),
Western Hemlock/Vanilla Leaf (23 subplots),
Western Hemlock/Oregon Grape (two subplots),
Western Hemlock/Oregon Grape-Salal (20 sub-
plots), and Western Hemlock/Salal (11 subplots)
(Figure 4). This diversity of habitat types at the
subplot level reflects the diversity of microhabitat
across the site. The wetter north and northeast
portions19 of the plot keyed to wetter plant associ-
ations characterized by lady fern, foamflower, and
vanilla leaf. The uphill portion of the plot is
characterized by salal and Oregon grape.
FOREST TREE CHARACTERIZATION
In the WRCCRF 4-ha plot directly under and
around the canopy crane, all trees 5 cm or larger
have been tallied, measured, and mapped (Table 3
Figure 4. Plant associations of the Wind River Canopy
Crane Research Facility 4-ha crane plot. Each square is
25 ? 25 m. The circle represents the swing of the crane jib
(2.3 ha), whereas the black square, just off center, is the
crane tower location. An ephemeral stream runs through
the northern portion of the plot, and an old logging road
(about 4 m wide) runs through the plot. Plant associa-
tions, from wet-site to dry-site indicators, are TSHE/ATFI,
Tsuga heterophylla/Athyrium filix-femina (Western Hem-
lock/Lady Fern) (2 subplots); TSHE/TITR, T. heterophylla/
Tiarella trifoliata (Western Hemlock/Foamflower) (4
subplots); TSHE/VAAL-GASH, T. heterophylla/Vaccinium
alaskaense-Gaultheria shallon (Western Hemlock/Alaska
Huckleberry-salal) (2 subplots); TSHE/ACTR, T. hetero-
phylla/Achlys triphylla (Western Hemlock/Vanilla Leaf)
(23 subplots); TSHE/BENE, T. heterophylla/Berberis nervosa
(Western Hemlock/Oregon Grape) (2 subplots); TSHE/
BENE-GASH, T. heterophylla/B. nervosa/G. shallon (West-
ern Hemlock/Oregon Grape-Salal) (20 subplots), and
TSHE/GASH, T. heterophylla/G. shallon (Western Hem-
lock/Salal) (11 subplots).
432 D. C. Shaw and others
Table 2. Vascular Plant Species in the Understory of the Wind River Old-growth Forest
Species/Form Scientific Name Family Ave. % Cover Frequency PV
Shrubs
Vine maple Acer circinatum Aceraceae 26.7 95 25.5
Salal Gaultheria shallon Ericaceae 16.0 100 16.0
Oregon grape Berberis nervosa Berberidaceae 14.2 100 14.2
Red huckleberry Vaccinium parvifolium Ericaceae 4.4 97 4.2
Baldhip rose Rosa gymnocarpa Rosaceae 0.7 55 0.4
Alaska V. alaskaense Ericaceae 0.6 31 0.2
huckleberry
Beaked hazelnut Corylus cornuta Betulaceae 0.4 28 0.1
Cascara Rhamnus purshiana Rhamnaceae 0.2 16 0.03
Snowberry Symphoricarpos albus Caprifoliaceae 0.1 8 0.01
Black V. membranaceum Ericaceae 0.1 6 0
huckleberry
Blackcap Rubus leucodermis Rosaceae 0.03 2 0
Honeysuckle Lonicera ciliata Caprifoliaceae 0.02 2 0
Salmonberry R. spectabilis Rosaceae 0.02 2 0
Herbs
Vanilla leaf Achlys triphylla Berberidaceae 4.4 100 4.4
Inside-out flower Vancouveria hexandra Berberidaceae 1.6 88 1.4
Twinflower Linnaea borealis Caprifoliaceae 1.3 88 1.1
Bear grass Xerophyllum tenax Liliaceae 1.4 73 1.0
Trillium Trillium ovatum Liliaceae 1.0 95 0.9
Queen cup bead Clintonia uniflora Liliaceae 0.9 88 0.8
lily
Trailing yellow Viola sempervirens Violaceae 0.9 86 0.8
violet
Rattlesnake Goodyera oblongifolia Orchidaceae 0.7 70 0.5
plantain
Hooker?s fairybell Disporum hookeri Liliaceae 0.6 61 0.4
Three-leaved Anemone deltoidea Ranunculaceae 0.6 59 0.4
Anemone35
Foamflower Tiarella trifoliata Saxifragaceae 0.6 53 0.3
Starry Smilacena stellata Liliaceae 0.6 53 0.3
Solomen pluume
Prince?s pine Chimaphila umbellata Pyrolaceae 0.5 52 0.3
False lily of the Maianthemum dilatatum Liliaceae 0.3 30 0.1
valley
Trailing Rubus ursinus Rosaceae 0.3 30 0.1
blackberry
Starflower Trientalis latifolia Primulaceae 0.3 30 0.1
Pathfinder Adenocaulon bicolor Asteraceae 0.3 27 0.1
Bedstraw Galium triflorum Rubiaceae 0.2 20 0.04
Bunchberry Cornus canadensis Cornaceae 0.2 16 0.03
dogwood
Twisted stalk Streptopus amplexifolius Liliaceae 0.2 16 0.02
Baneberry Actaea rubra Ranunculaceae 0.1 11 0.01
Menzies? Chimaphila menziesii Pyrolaceae 0.05 5 0
pipissewa
Dogbane Apocynum Apocynaceae 0.03 3 0
Androsaemifolium
Wild ginger Asarum caudatum Aristolochiaceae 0.03 3 0
Hairbell Campanula scouleri Campanulaceae 0.03 3 0
Hawksbeard Hieracium albiflorum Asteraceae 0.03 3 0
Veronica Veronica americana Scrophulariaceae 0.02 2 0
Table 2. Continued
Wind River 2Old-growth Forest 433
and Figure 5). The forest has eight coniferous spe-
cies, including Douglas-fir, western hemlock,
western red cedar (Thuja plicata), Pacific yew (Taxus
brevifolia), Pacific silver fir, noble fir (Abies procera),
grand fir (Abies grandis), western white pine (Pinus
monticola), and two small stature angiosperms,
cascara (Rhamnus purshiana) and Pacific dogwood
(Cornus nuttallii). Red alder (Alnus rubra) occurs in
some canopy gaps along the ephemeral stream.
Here, the 1999 basal area of conifers is 82.9 m2
ha)1, falling in the midrange (50?129 m2 ha)1) of
450- to 500-year-old stands reported by Franklin
and Waring (1980) in the Pacific Northwest. Av-
erage density of all trees is 427 stems ha)1. The
majority of trees are western hemlock (224.0 trees
ha)1); however, Douglas-fir is the dominant species
with respect to basal area (35.4 m2 ha)1), followed
by western hemlock (26.9 m2 ha)1) and western
red cedar (16.5 m2 ha)1). Average height of
Douglas-fir is 52.0 m (Figure 5), with the tallest
tree being 64.6 m, much less than the maximum
heights of Douglas-fir, which often reach 70?80 m
in more favorable sites (Franklin and Waring
1980). Western hemlock height averages 19.0 m,
with the tallest tree being 55.4 m. This is in the
range of maximum attainable heights (50?65 m)
reported by Franklin and Waring (1980). The ver-
tical structure of the forest (Figure 1) is described in
detail by Parker and colleagues (2004). The height
and diameter distribution of Douglas-fir indicate20 a
classic shade-intolerant cohort of trees that are not
being replaced by reproduction (Figure 5). Western
hemlock, western red cedar, Pacific yew, and Pa-
cific silver fir are shade tolerant and reproducing
well (Figure 5).
DISTURBANCE AND DYNAMICS
The primary disturbance regime for Douglas-fir
forests in this region is catastrophic, stand-re-
placement (high to moderate severity) fire, which
appears to have occurred in the past at return in-
tervals of 300?650 years (Hemstrom and Franklin
1982; Agee 1991; Gray and Franklin 1997). The
Wind River old-growth forest most likely originated
after a high-severity fire, or series of fires, about
500 years before present. The Douglas-fir trees,
estimated to be 375?500 years old (J. Franklin
personal communication), were dated by counting
growth rings on stumps surrounding the T. T.
Munger RNA. We have cored 17 large-diameter
western hemlock trees within the old-growth
Table 2. Continued
Species/Form Scientific Name Family Ave. % Cover Frequency PV
Ferns
Bracken36 Pteridium aquilinum Polypodiaceae 1.5 91 1.4
fern
Deer fern Blechnum spicant Polypodiaceae 2.1 38 0.8
Sword fern Polysticum munitum Polypodiaceae 0.6 42 0.3
Lady fern Athyrium felix-femina Polypodiaceae 0.5 25 0.1
Sierra wood fern Thelypteris nevadensis Polypodiaceae 0.4 19 0.1
Running Lycopodium clavatum Lycopodiaceae 0.1 8 0.01
clubmoss
Graminoids
Woodrush Luzula campestris Juncaceae 0.03 3 0
Nodding trisetum Trisetum cernuum Poaceae 0.02 2 0
Nonchlorophyllous ericad?s
Indian pipe Monotropa uniflora Ericaceae 0.05 5 0
Candystick Allotropa virgata Ericaceae 0.02 2 0
Pinesap Hypopitys monotropa Ericaceae 0.02 2 0
Pinedrops Pterospora andromedea Ericaceae 0.02 2 0
Nonnative weeds
Plantain Plantago major Plantaginaceae 0.1 6 0
Self-heal Prunella vulgaris Labiatae 0.05 5 0
Pearly everlasting Anaphalis margaritacea Asteraceae 0.02 2 0
White clover Trifolium repens Fabaceae 0.02 2 0
Festuca Festuca pratensis Poaceae 0.02 2 0
Data are based on sixty-four 25 ? 25-m subplots that make up the Wind River Canopy Crane Research Facility 4-ha plot. Ave. % Cover is the mean cover estimated on all 64
subplots. Frequency is the percentage37 of subplots a plant species occurred in. PV, prominence value (Ave. % cover * frequency).
434 D. C. Shaw and others
stand, and none of them are over 300 years of age.
Douglas-fir probably dominated the young forest
for the first 200 years, after which the shade-tol-
erant western hemlock and western red cedar were
recruited into the understory.
A series of mortality strips and growth plots were
installed in the T. T. Munger RNA in 1947 (King
1961) and provide information on the recent dis-
turbance dynamics of this 500-year-old forest. De-
Bell and Franklin (1987) and Franklin and DeBell
(1988) have analyzed 36 years of this record
(1947?83, collected in 6-year intervals) for tree
population change, growth, and mortality. The
annual rate of mortality for all trees during this
period was 0.75%, with 22% of the original stems
dying. Western hemlock, western red cedar, and
Pacific silver fir, all shade-tolerant species, were
recruited into the stand so that the total number of
live trees ha)1 was maintained (1947 = 479.5 tree
ha)1; 1983 = 443.6 trees ha)1). Douglas-fir ac-
counted for 33% of the volume (m3 ha)1) growth
yet nearly 50% of the mortality volume. Western
hemlock accounted for 50% of the volume growth
but only 28% of the mortality volume. Pacific silver
fir increased in the lower canopy, accounting for
60% of the small tree ingrowth. The Douglas-fir
trees are slowly dying out of the stand because
there is no reproduction of this shade-intolerant
species in the understory. The extinction of Doug-
las-fir is predicted in 740 years at the current rate of
mortality (Franklin and DeBell 1988). Four canopy
processes are responsible for the long-term survival
of individual Douglas-fir trees (Ishii and Ford 2002;
Ishii and others 2000a, 2000b). These include ver-
tical stratification (Douglas-fir dominates the upper
canopy), decreasing crown competition in the up-
per canopy, morphological acclimation (old Doug-
las-fir have a unique crown form and branching
pattern that maintains efficient shoot and foliage
display), and crown maintenance (epicormic
branching at the twig, branch, and stem levels
maintains foliage).
The 125-year age spread of the Douglas-fir trees is
typical for stand initiation in the region. One reason
may be reburns of areas that were originally cata-
strophically burned (Gray and Franklin 1997),
although competition with shrubs and seed source
may also play a role. Although reburns may occur,
light ground fires within forests are rare, and there is
no evidence in the T. T. Munger RNA that, once the
forest stand was fully established, fire has occurred.
Wind disturbance is common in the region, but
catastrophic wind events are associated with forests
on exposed westerly slopes and ridgetops whereas
the Wind River old-growth forest is in a north?
south-oriented valley bottom, protected from the
most severe wind effects coming from the coast.
Mortality in this forest is associated with un-
known causes (30%), windthrow (23%), broken
bole or top (22%), mechanical snow damage
(10%), crushing (7%), and insects and diseases
(6%) (Franklin and DeBell 1988). Windthrow
events tend to be at the individual tree level and
associated with root-rot and butt-rot fungi for up-
rooting and butt snap, as well as heart-rot fungi for
stem breakage (Bible 2001). An episode of Douglas-
fir bark beetle (Dendroctonus pseudotsugae)-caused
mortality was reported for the 1950s, and Franklin
Table 3. Conifer Tree Speciesa in the Wind River Old-growth Forest
Tree species Trees ha)1
Standing
Dead ha)1
DBHb (cm)
(mean)
Height
(m) (mean)
Basal area
m2 ha)1
Abies amabilis 47.0 9.2 12.2 8.5 1.2
Abies grandis 4.5 5.0 51.3 39.1 1.0
Abies procera 0.5 0.0 49.8 32.3 0.1
Pinus monticola 0.5 4.0 48.0 32.1 0.1
Pseudotsuga 35.0 54.5 111.0 52.0 35.4
menziesii
Taxus brevifolia 85.8 15.8 13.8 6.2 1.7
Thuja plicata 30.0 3.5 68.6 29.0 16.5
Tsuga 224.0 18.0 29.9 19.0 26.9
heterophylla
Total conifers 427.3 110.0 34.2 19.2 82.9
Data are ha)1, averaged from the Wind River Canopy Crane Research Facility 4-ha plot. Standing dead tree minimum height, 1.37 m. Three angiosperm tree species (Cornus
nuttalii,Alnus rubra, and Rhamnus purshiana) are also present, but not abundant.
aWith DBH greater than 5 cm.
bDBH, diameter at breast height.
Wind River 2Old-growth Forest 435
and DeBell feel that much of the unknown
mortality of Douglas-fir may be associated with this
beetle, in conjunction with root disease and other
stressors such as drought. White pine blister rust
(Cronartium ribicola), a nonnative disease, is asso-
ciated with decline and mortality of western white
pine, although mountain pine beetle (Dendroctonus
ponderosae) may also play a role. Hemlock dwarf
mistletoe (Arceuthobium tsugense) is common in the
T. T. Munger RNA and may weaken and kill
western hemlock (DeBell and Franklin 1987). It
occurs over about 1 ha of the WRCCRF 4-ha plot in
a distinct infection center (Mathiasen and Shaw
1998; Shaw and others 2000). Pacific silver fir
mortality is associated with Armillaria root disease
(Armillaria spp.), which is especially virulent on
silver fir at the lower limits of the Pacific silver fir
vegetation zone (G. MacDonald personal commu-
nication).
Herbivory 21(folivory) is estimated at less than
2% y)1 for the canopy trees at the canopy crane
site (Schowalter and Ganio 1998; Shaw and oth-
ers unpublished 22). Defoliator outbreaks in old-
growth Douglas-fir forests west of the Cascade
crest are rare (Perry and Pitman 1983). Herbivory
at Wind River is much greater in the understory,
where an annual rate of 9.9% was documented
for vine maple in 1999 (Braun and others 2002).
Deer and elk also play a significant role in un-
derstory herbivory. We have observed consid-
erable browsing by these animals on salal,
huckleberries, vine maple, and bear grass, as well
as small Pacific yew and western red cedar.
OLD-GROWTH FORESTS
Characterizing a forest as ??old growth?? is
important because of the theoretical and practical
concepts concerning how stand age and produc-
tivity will influence carbon dynamics. Tree pro-
ductivity, forest-stand productivity, and live-tree
biomass accumulation all decline with age (Bond
and Franklin 2002). Decreasing net primary pro-
ductivity and increasing mortality account for the
decline in live-tree biomass accumulation in
Douglas-fir forests (Acker and others 2002). How-
ever, carbon dynamics also include total forest car-
bon accumulation (sequestration), which increases
with stand age in the Pacific Northwest of North
America (Janisch and Harmon 2002). Whether old-
growth forests will function as a carbon sink or
source to the atmosphere is a critical question asked
in this special feature (Paw U and others 2004).
Predicting carbon storage and exchange with the
atmosphere at the landscape level requires scaling
the various age classes and management regimes
(Harmon 2001), and23 an important consideration
includes the extent and distribution of old growth.
Old-growth forests may currently occur on about
4.12 million ha in Washington, Oregon, and
northern California (Bolsinger and Waddell 1991),
although definitions vary and not all experts agree
on this number (Margot and others 1991). This may
be about 17% of the original old growth that
occupied the region in the mid 1800s, although this
is also debated (Margot and others 1991) and may
by closer to 10%.
Old-growth Douglas-fir forests are characterized
by large, live old (175?350 years +) trees, a deep
(50 m +) vertically continuous canopy, large snags,
large logs on land, and large logs in streams
(Franklin and others 1981; Franklin and Spies
1991). Old-growth forests retain nutrients; accu-
mulate organic matter; reduce peak flows during
rainfall runoff; are heterogeneous; provide opti-
mum habitat for specialized vertebrates; have well-
Figure 5. Tree diameter [10-cm diameter at breast
height (DBH) classes] and height (10-m height classes)
class distribution of major conifer tree species on the
Wind River Canopy Crane Research Facility 4-ha crane
plot. Data for trees is the average ha)1.
436 D. C. Shaw and others
developed understory vegetation; have a high di-
versity of arthropods, fungi, lichens, and bryo-
phytes; and maintain high gross productivity,
although mortality usually balances growth
(Franklin and others 1981). A unique characteristic
of Pacific Northwest old-growth forests is the long
persistence of primary successional species, such as
Douglas-fir and western white pine (Pinus montico-
la), which can live for 400?1000 years (Franklin
1988).
Structural features that define a forest as an old-
growth Douglas-fir forest in western Washington
and Oregon, on western hemlock sites, are 10?20
Douglas-fir trees larger than 80-cm diameter at
breast height (DBH) ha)1, 20 trees of any species
larger than 80-cm DBH ha)1, 10?30 shade-tolerant
trees larger than 40-cm DBH ha)1, 4?10 snags
larger than 50-cm DBH and larger than 5 m tall
ha)1, and a minimum log biomass of 30?34 tons
ha)1 (Franklin and Spies 1991). The WRCCRF 4-ha
plot averages 31.5 Douglas-fir ha)1 larger than 81-
cm DBH, 78.5 shade-tolerant-associated tree spe-
cies (western hemlock, western red cedar, Pacific
silver fir) ha)1 larger than 41-cm DBH, 59.5 snags
ha)1 larger than 51-cm DBH (although we did not
include minimum height of 5 m), and log biomass
is 93.9 Mg ha)1. Therefore, the Wind River forest
fits the structure-based definition of an old-growth
forest.
GLOBAL CONTEXT
The International Biological Program, Wood-
lands Data Set (DeAngelis and others 1981), rep-
resents the range of mean annual rainfall and
temperature conditions for forests throughout the
globe. The Wind River old-growth forest lies at an
extreme combination of cool and wet for sites
summarized in this dataset (Figure 6). There are
other colder forested sites but they are drier, and
there are many warmer forested sites but they can
be wetter. Structurally, the stand has one of the
higher aboveground biomass numbers reported
globally, although it is at the low end of the range
for old-growth forests in the Pacific Northwest
(Harmon and others 2004). The forest is charac-
terized as a temperate coniferous seasonal rain
forest due to the high rainfall (2223 mm y)1), lat-
itudinal position (N 45
), and the extreme season-
ality of the rainfall (less than 10% during summer)
(Schoonmaker and others 1997).
ACKNOWLEDGEMENTS
The Wind River Canopy Crane Research Facility is
a cooperative scientific venture between the Uni-
versity of Washington, College of Forest Resources,
USDA Forest Service Pacific Northwest Research
Station, and the Gifford Pinchot National Forest.
Joel Norgren and Ted Dyrness provided the soil
profile descriptions of the T. T. Munger RNA and
input on the soils section. Tom High also provided
input on the soils section. Some of this work was
supported by the Office of Science, Biological and
Environmental Research Program (BER), US De-
partment of Energy (DOE), through the Western
Regional Center (WESTGEC) of the National In-
stitute for Global Environmental Change (NIGEC)
under Cooperative Agreement DE-FCO3-
90ER61010. Any opinions, findings, and conclu-
sions or recommendations expressed herein are
those of the authors and do not necessarily reflect
the view of DOE.
REFERENCES
Acker SA, Halpern CB, Harmon ME, Dyrness CT. 2002. Trends in
bole biomass accumulation, net primary production and tree
mortality in Pseudotsuga menziesii forests of contrasting age.
Tree Physiol 22:213?7.
Figure 6. Mean temperature and precipitation for a
global distribution of representative forests (circles) from
the International Biological Program, Woodlands Dataset
(DeAngelis and others 1981) (ORNL DAAC archive at
www.ornl.gov). The Wind River Canopy Crane Research
Facility (WRCCRF) site is added to this figure, located in
the top left-hand corner as a solid square, to provide a
reference for the global climatic regime of the site. The
Wind River old-growth forest is at the extreme combi-
nation of cool and wet for sites summarized in this da-
taset.
Wind River 2Old-growth Forest 437
Agee JK. 1991. Fire history of Douglas-fir forests in the Pacific
Northwest. In: Ruggiero LF, Aubry KB, Carey AB, Huff MH,
technical coordinators. Wildlife and vegetation of unmanaged
Douglas-fir forests. General technical report PNW-GTR-285.
Portland (OR): USDA Forest Service, Pacific Northwest Re-
search Station. p 25?33.
Bible KJ. 2001. Long-term patterns of Douglas-fir and western
hemlock mortality in the Cascade Mountains of Oregon and
Washington [dissertation]. Seattle: University of Washington24 .
Bolsinger CL, Waddell KL. 1993. Area of old-growth forests in
California, Oregon and Washington. Resource bulletin PNW-
RB-197. Portland (OR):25 USDA Forest Service, Pacific North-
west Research Station.
Bond BJ, Franklin JF. 2002. Aging in Pacific Northwest forests: a
selection of recent research. Tree Physiol 22:73?6.
Brady NC, Weil RR. 1996. The nature and property of soils.
Prentice Hall: EnglewoodCliffs (NJ).
Brockway DG, Topik C, Hemstrom MA, Emmingham WH. 1983.
Plant association and management guide for the Pacific Silver
Fir Zone, Gifford Pinchot National Forest. R6-Ecol-130a-1983.
Portland (OR): USDA Forest Service, Pacific Northwest Re-
search Station.
Braun DM, Runcheng B, Shaw DC, Van Scoy M. 2002. Folivory
of vine maple in an old-growth Douglas-fir?western hemlock
forest. Northwest Sci 76:315?21.
Chen J, Franklin JF, Spies TA. 1992. Vegetation responses to
edge environments in old-growth Douglas-fir forests. Ecol
Appl 2:387?96.
Chen J, Franklin JF, Spies TA. 1993a. An empirical model for
predicting diurnal air-temperature gradients from clearcut-
forest edge into old-growth Douglas-fir forest. Ecol Modell
67:179?98.
Chen J, Franklin JF, Spies TA. 1993b. Contrasting microclimate
patterns among clearcut, edge, and interior area of old-growth
Douglas-fir forest. Agric For Meteorol 63:219?37.
Cromack K Jr, Miller RE, Helgerson OT, Smith RB, Anderson
HW. 1999. Soil carbon and nutrients in a coastal Douglas-fir
plantation with red alder. Soil Sci Soc Am J 63:232?9.
DeAngelis DL, Gardner RH, Shugart HH. 1981. Productivity of
forest ecosystems studied during the IBP: the woodlands data
set. In: Reichle DE editor. Dynamic properties of forests ec-
osystems. Cambridge: Cambridge University Press. p 567?
673.
DeBell DS, Franklin JF. 1987. Old-growth Douglas-fir and
western hemlock: a 36-year record of growth and mortality.
West J Appl For 2:111?4.
Franklin JF. 1972. Wind River Research Natural Area.Federal
Research Natural Areas in Oregon and Washington: a guide-
book for scientists and educators.26 Portland: Pacific Northwest
Forest and Range Experiment Station.USDA Forest Service
WR-1 p?WR-12.
Franklin JF. 1988. Pacific Northwest forests. In: Barbour MG,
Billings WD editors.. North American terrestrial vegetation.
Cambridge: Cambridge University Press. p 103?30.
Franklin JF, Cromack K Jr, Denison W, McKee A, Maser C,
Sedell J, Swanson F, Juday G. 1981. 16. Ecological charac-
teristics of old-growth Douglas-fir forests. General technical
report PNW-118. Portland (OR): USDA Forest Service, Pacific
Northwest Research Station.
Franklin JF, DeBell DS. 1988. Thirty-six years of tree popula-
tions change in an old-growth Pseudotsuga? Tsuga forest. Can J
For Res 18:633?9.
Franklin JF, Dyrness CT. 1973. Natural vegetation of Oregon and
Washington. General technical report PNW-8. Portland (OR):
USDA Forest Service, Pacific Northwest Research Station.
417 p.
Franklin JF, Spies TA. 1991. Ecological definitions of old-growth
Douglas-fir forests. In: Ruggiero LF, Aubry KB, Carey AB, Huff
MH, technical coordinators. Wildlife and vegetation of un-
managed Douglas-fir forests. General technical report PNW-
GTR-285. Portland (OR): USDA Forest Service, Pacific
Northwest Research Station. p 71?80.
Franklin JF, Waring RH. 1980. Distinctive features of the
northwestern coniferous forest: development, structure, and
function. In: Ecosystem analysis: proceedings, 40th Annual
Biological Colloquium, 1979 April 27?28. Corvallis: Oregon
State University. p 59?8627 .
Gray AN, Franklin JF. 1997. Effects of multiple fires on the
structure of southwestern Washington forests. Northwest Sci
71:174?85.
Hammond PE. 1980. Reconnaissance geologic map and cross
sections of southern Washington Cascade Range, latitude 45
degrees 30 minutes?47 degrees 15 minutes N., longitude 120
degrees 45 minutes?122 degrees 22.5 minutes W. Portland:
Portland State University.
Hammond PE, Korosec MA. 1983. Geochemical analyses, age
dates, and flow-volume estimates for Quaternary volcanic
rocks, southern Cascade Mountains, Washington. Open
file report 83-13. Olympia (WA): Washington Department
of Natural Resources, Division of Geology and Earth Resouces.
p 36.
Harmon ME. 2001. Carbon sequestration in forests. J For 99:
24?9.
Harmon ME, Bible K, Ryan MG, Shaw DC, Chen H, Klopatek J,
Li X. 2004. Production, respiration, and overall carbon balance
in an old-growth Pseudotsuga? Tsuga forest ecosystem. Eco-
systems 7:498?51228 .
Harr RD. 1986. Effects of clearcutting on rain-on-snow runoff in
western Oregon: A new look at old studies. Water Resour Res
22:1095?100.
Heilman PE. 1981. Minerals, chemical properties and fertility of
forest soils. In: Heilman PE, Anderson HW, Baumgartner DM
editors.. Forest soils of the Douglas-fir region. Pullman:
Washington State University. p 121?36.
Hemstrom MA, Franklin JF. 1982. Fire and other disturbances of
the forests in Mount Rainier National Park. Quat Res 18:32?51.
Homan PS, Lollins P, Chappell HN, Stangenberger AG. 1995. Soil
organic carbon in a mountainous, forested region: relation to
site characteristics. Soil Sci Soc Am 59:1468?75.
Ishii H, Ford ED. 2002. Persistance of Pseudotsuga-menziesii in
temperate forests of the Pacific Northwest Coast, USA. Folia
Geobotanica 37:63-929 .
Ishii H, Clement JP, Shaw DC. 2000a. Branch growth and
crown form in old coastal Douglas-fir. For Ecol Manage 131:
81?91.
Ishii H, Reynolds JH, Ford ED, Shaw DC. 2000b. Height growth
and vertical development of an old-growth Pseudotsuga? Tsuga
forest in southwestern Washington, U.S.A. Can J For Res
30:17?24.
Janisch JE, Harmon ME. 2002. Successional changes in live and
dead wood carbon stores: implications for net ecosystem
productivity. Tree Physiol 22:77?89.
Kemp L, Schuller SR. 1982. Checklist of vascular plants of the
Thornton T. Munger Research Natural Area. Administrative
438 D. C. Shaw and others
report PNW-4. Portland (OR). USDA Forest Service. Pacific
Northwest Forest and Range Experiment Station.
King JP. 1961. Growth and mortality in the Wind River Natural
Area. J For 59:768?70.
Lawrence DB. 1939. Some features of the vegetation of the
Columbia River Gorge with special reference to asymmetry in
forest trees. Ecol Monogr 9:217?57.
Margot BG, Holthausen RS, Teply J, Carrier WD. 1991. Old-
growth inventories: status, definitions, and visions for the
future. In: Ruggiero LF, Aubry KB, Carey AB, Huff MH,
technical coordinators. Wildlife and vegetation of unmanaged
Douglas-fir Forests. General technical report PNW-GTR-285.
Portland (OR): USDA Forest Service, Pacific Northwest
Research Station. p 47?60.
Mathiasen RL, Shaw DS. 1998. Adult sex ratio of western
hemlock dwarf mistletoe in six heavily infected western
hemlock. Madrono 45:210?4.
Means JE, MacMillan PC, Cromack K Jr. 1992. Biomass and
nutrient content of Douglas-fir logs and other detrital pools in
an old-growth forest, Oregon, U.S.A. Can J For Res 22:1536?
46.
Meyers AP, Fredricks N. 1993. Thornton T. Munger Research
Natural Area management plan. USDA Forest Service. On file
Gifford Pinchot National Forest. Vancouver (WA). p 62.
Parker GG, Harmon ME, Lefsky MA, Chen J, Van Pelt R, Weiss
SB, Thomas SC, Winner WE, Shaw DC, Franklin JF. 2004.
Three-dimensional structure of an old-growth Pseudotsuga?
Tsuga canopy and its implications for radiation balance,
microclimate, and atmospheric gas exchange Ecosystems
30 7:440?53.
Paw U KT, Falk M, Suchanek TH, Ustin SL, Chen J, Park Y-S,
Winner WE, Thomas SC, Hsiao TC, Shaw RH, and others.
2004. Carbon dioxide exchange between an old-growth forest
and the atmosphere. Ecosystems31 7:513?24.
Perry DA, Pitman GB. 1983. Genetic and environmental influ-
ences in host resistance to herbivory: Douglas-fir and the
western spruce budworm. Z Angew Entomol 96:217?28.
Radwan MA. 1992. Effect of forest floor on growth and nutrition
of Douglas-fir and western hemlock seedlings with and
without fertilization. Can J For Res 22:1222?9.
Schowalter TD, Ganio LM. 1998. Vertical and seasonal variation
in canopy arthropod communities in an old-growth conifer
forest in southwestern Washington, USA. Bull Entomol Res
88:633?40.
Schoonmaker PK, Von Hagen B, Wolf EC. 1997. The rain forests
of home. Washington (DC): Island.
Shaw DC, Freeman EA, Mathiasen RL. 2000. Evaluating the
accuracy of ground based hemlock dwarf mistletoe rating: a
case study using the Wind River canopy crane. West J Appl
For 15:8?14.
Thomas SC, Winner WE. 2000. Leaf area index of an old-growth
Douglas-fir forest: an estimate based on direct structural
measurements in the canopy. Can J For Res 30:1922?30.
Topik C, Halverson NM, Brockway DG. 1986. Plant association
and management guide for the Western Hemlock Zone, Gifford
Pinchot National Forest. R6-ECOL-230A-1986. Portland (OR):
USDA ForestService, Pacific Northwest Research Station.p 133.
Walsh TJ, Korosec MA, Phillips WM, Logan RL, Schasse HW.
1987. Geologic map of Washington: southwest quad-
rant?geologic map GM-34. Olympia: Washington State
Department of Natural Resources.
Wise WS. 1961. Geology and mineralogy of the Wind River area,
Washington, and the stability relations of celadonite [disser-
tation]. Baltimore (MD): Johns Hopkins University32 .
Wise WS. 1970. Cenozoic volcanism in the Cascade Mountains
of southern Washington. Wash Div Mines Geol Bull 60:1?45.
Woodfin RO Jr, DeBell DS, Franklin JF. 1987. Wind River Ex-
perimental Forest Research Management Plan. USDA Forest
Service. On file: Pacific Northwest Forest and Range Experi-
ment Station (now, Pacific Northwest Research Station),
Portland (OR).
Youngberg CT. 1966. Forest floor in Douglas-fir forests. I.
Dry weight and chemical properties. Soil Sci Soc Am J 30:
406?9.
Wind River 2Old-growth Forest 439