FEIS Home Page |
AUTHORSHIP AND CITATION:
Gucker, Corey L. 2005. Cornus nuttallii.
In: Fire Effects Information System, [Online].
U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Laboratory (Producer).
Available: https://www.fs.usda.gov/database/feis/plants/tree/cornut/all.html [].
FEIS ABBREVIATION:
CORNUT
SYNONYMS:
None
NRCS PLANT CODE [128]:
CONU4
COMMON NAMES:
Pacific dogwood
mountain dogwood
mountain flowering dogwood
TAXONOMY:
The scientific name of Pacific dogwood is Cornus nuttallii Audubon ex.
Torr. and Gray (Cornaceae) [60,63,64,72].
Pacific dogwood, C. florida, and C. kousa are sexually compatible. These hybrids are commercially recognized [112]:
C. nuttallii × florida
(C. florida × C. kousa) × C. nuttallii
When literature is cited in this review that refers to the Cornus genus
only, it will be indicated as Cornus spp.
LIFE FORM:
Tree-shrub
FEDERAL LEGAL STATUS:
None
OTHER STATUS:
Pacific dogwood is listed as a priority 1 species in the state of Idaho [68,69].
This listing states the "taxa is in danger of becoming extinct or extirpated
from Idaho in the foreseeable future if identifiable factors contributing to their
decline continue to operate..."
The Flora of North America
provides a distributional map of Pacific dogwood.
ECOSYSTEMS [45]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES27 Redwood
FRES28 Western hardwoods
STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES
CA | ID | OR | WA |
BC |
Washington: Cascade Range: In Douglas-fir-western hemlock-Pacific silver fir (Abies amabilis) mixed forests, pacific dogwood is a common subcanopy species. Other canopy vegetation can include grand fir (A. grandis), noble fir (A. procera), Pacific yew (Taxus brevifolia), western redcedar (Thuja plicata), and bigleaf maple (Acer macrophyllum). Associated shrub species are vine maple (A. circinatum), Oregon-grape (Mahonia repens), salal (Gaultheria shallon), red huckleberry (Vaccinium parvifolium), and Pacific rhododendron (Rhododendron macrophyllum) [17,29,43]. In Mount Rainier National Park, Pacific dogwood occurs in western hemlock/sweet after death (Achlys triphylla) and western hemlock/salal vegetation associations [44].
Oregon: Cascade Range: In the west-central Cascades, Pacific dogwood occurs in old-growth Douglas-fir forests. Pacific dogwood coverage is greatest in Douglas-fir/Pacific rhododendron communities that occupy warm, mesic, south slopes. Coverage of Pacific dogwood is less in Douglas-fir/giant chinquapin (Chrysolepis chrysophylla) and Douglas-fir/Pacific rhododendron/Cascade barberry (M. nervosa) communities that are found on xeric south slopes and on mesic sites, respectively [49]. Pacific dogwood is also typical in Douglas-fir/vine maple/western sword fern (Polystichum munitum), western hemlock/vine maple/western sword fern, and western hemlock/salal communities [110].
Coast Range: Along the east slope of the Coast Range, Pacific dogwood occurs in both the vine maple/salal and oceanspray (Holodiscus discolor)/salal community types. Coverage is normally greater in the oceanspray/salal community [3]. In the Douglas-fir forests of the southern Coast Range, Pacific dogwood is commonly associated with California bay (Umbellularia californica), tanoak (Lithocarpus densiflora), and evergreen huckleberry (V. ovatum) [26]. On the eastern side of the Coast Range, Pacific dogwood in typically found with Douglas-fir, western redcedar, bigleaf maple, Oregon ash (Fraxinus latifolia), and bitter cherry (Prunus emarginata) [28,50].
Siskiyou Region: Pacific dogwood is common in tanoak-California bay/Pacific rhododendron and tanoak-redwood (Sequoia sempervirens) vegetation types of southwestern Oregon [13].
Willamette Valley: At low elevations in the Willamette Valley, Pacific dogwood occurs in Oregon white oak (Quercus garryana) forests. Other associated vegetation includes, California black oak (Q. kelloggii), canyon live oak (Q. chrysolepis), bigleaf maple, Oregon ash, ponderosa pine (Pinus ponderosa), incense-cedar (Calocedrus decurrens), Pacific madrone (Arbutus menziesii), and tanoak [120,125].
Oregon/California: Coast Range: In the Douglas-fir-tanoak-Pacific madrone cover type, Pacific dogwood is found with salal, evergreen huckleberry, Oregon-grape, Pacific rhododendron, and poison-oak (Toxicodendron diversilobum) [113]. In redwood forests of Oregon and California, Douglas-fir, western hemlock, tanoak, California hazel, salal, Pacific ninebark (Physocarpus capitatus), and Pacific rhododendron commonly occur with Pacific dogwood [107].
Klamath Range: Within the Pacific ponderosa pine (Pinus ponderosa var. ponderosa) -Douglas-fir forest type of the Klamath Mountains, Pacific dogwood is common [86]. Pacific dogwood is also in the understory of white fir (Abies concolor)/dwarf Oregon-grape and mixed evergreen forests. More unique species in these forests include, sugar pine (P. lambertiana), Port-Orford-cedar (Chamaecyparis lawsoniana), and California hazel (Corylus cornuta var. californica) [114,134].
California: Coast Range: Pacific dogwood is common in Douglas-fir-hardwood and ponderosa pine forests of the California Coast Range [18,20,115]. In ponderosa pine forests, typical overstory species are sugar pine, Douglas-fir, gray pine (P. sabiniana), white fir, incense cedar, and Oregon white oak. Understory associates include manzanitas (Arctostaphylos spp.), deerbrush (Ceanothus integerrimus), California coffeeberry (Rhamnus californica), birchleaf mountain-mahogany (Cercocarpus betuloides), common snowberry (Symphoricarpos albus), and poison-oak [18,20].
Sierra Nevada Range: In the Sierra Nevada Range, Pacific dogwood is common in ponderosa pine/mixed conifer forests that include tanoak, California black oak, Sierra mountain misery (Chamaebatia foliolosa), and poison-oak [4]. Pacific dogwood is also typical of forests described as Sierra Nevada-Sierran montane and Sierra Nevada mixed conifer types [7,16,123]. In giant sequoia (Sequoiadendron giganteum) forests, Pacific dogwood, canyon live oak, Scouler willow (Salix scouleriana), white alder (Alnus rhombifolia), California hazel, whiteleaf manzanita (Arctostaphylos viscida), Sierra Mountain misery, California wildrose (Rosa californica), and Sierra gooseberry (Ribes roezlii) make up the understory vegetation [75,109].
Idaho: Pacific dogwood in the Lochsa-Selway area of northern Idaho is found in the western red cedar-western hemlock vegetation zone [105]. Hickey [59] reports that Pacific dogwood is also associated with Douglas-fir, grand fir, bitter cherry, oceanspray, Saskatoon serviceberry (Amelanchier alnifolia), common snowberry, Scouler willow, thimbleberry (Rubus parviflorus), Rocky Mountain maple (Acer glabrum) and red-osier dogwood (Cornus sericea).
|
|
GENERAL BOTANICAL CHARACTERISTICS:
This description provides
characteristics that may be relevant to fire ecology, and is not meant for
identification. Keys for identification are available [60,63,64,67,91,98].
Pacific dogwood is a native, deciduous, multi-branched tree, sometimes considered a shrub. Average mature-height estimates range from 20 to 75 feet (6.1-22.9 m) and canopy spread is often 20 feet (6.1 m) [60,64,76,83,85,104]. Young bark is thin and smooth, but ridges develop later making the trunk appear scale like [23,98]. The maximum trunk diameter reported for Pacific dogwood was 24 inches (61cm) [67]. The root system, commonly a taproot, penetrates deeply [67].
The growth form of Pacific dogwood may change with site conditions. When grown under a canopy of vegetation, the trunk is normally tapered and the crown is slender and short. When developed under a sparse canopy or in the open, the trunk is typically shorter, and the rounded crown can be as wide as it is tall [67]. Pacific dogwood branches have fine hairs and bear simple, opposite leaves that measure between 2 and 5 inches (5.1-12.7 cm) long by 1.5 to 2.8 inches (3.8-7.1cm) wide [11,60,64,67,91,98]. Leaves are hairier on the underside but have stiff appressed hairs above [60,91]. Pacific dogwood bears 2-seeded drupe fruits that are 1 to 1.5 cm long [63,64,91,98]. Commonly each drupe is comprised of 20-40 drupelets that are slightly flattened from being held in a tight cluster; contained seeds are smooth [67,91].
As a subcanopy species, Pacific dogwood has several shade growing adaptations. At 1/3 full sunlight, Pacific dogwood maintains maximum photosynthetic potential [11]. Branches are self-shading; leaf petioles orient downward allowing leaves to rest on and shade the branches. Although the trunk of Pacific dogwood can be damaged by direct sunlight [84,104], established plants may initiate shoot growth from the crown to shade and protect the exposed trunk [104].
Although typically considered a mesic species, Pacific dogwood is quite drought tolerant. The osmotic potential at zero turgor is -2.2 MPa; leaves begin to lose turgor pressure at 16-18% relative water deficit [108].
Botanical characteristics are altered when plants are infected with dogwood
anthracnose, a nonnative fungal disease caused by Discula spp.,
common in Pacific dogwood [22,24,30,31,32]. Fungal activity is usually greatest
from May through early July. However, the fungus can be active any time conditions
are moist and plants are growing [24]. Infected leaves develop blotches and
often drop early. Defoliation can be extreme. Twigs with this fungal disease are
depressed in spots and allow the fungus to progress into leaf buds, killing them
and setting back spring emergence [32]. Seed production diminishes with anthracnose
infection [38]. This fungal disease is considered threatening to native Pacific
dogwood populations because of its rapid spread and severe effects [25]. Control
measures have been described by many [24,30,31,32].
RAUNKIAER [Raunkiaer] LIFE FORM:
Phanerophyte
REGENERATION PROCESSES:
Pacific dogwood reproduces both sexually and asexually. Regeneration occurs
through seed production and/or by vegetative sprouting [3,82,104,105].
Breeding system: Outcrossing is likely common in Pacific dogwood as pollination is insect mediated [6]. Autogamy was not discussed in the literature.
Pollination: Pacific dogwood flowers are chiefly pollinated by insects [6].
Seed production: Reports of seed production by Pacific dogwood vary considerably. Lichthardt [82] suggests Pacific dogwood inflorescences produce 20 to 60 head flowers, each of which produces a single seed, but others consider Pacific dogwood drupes to be 2-seeded [63,64]. Brush [23] claims that Pacific dogwood produces abundant seed annually. Brinkman and Vankus [21], in a review, report that there are usually 2 years between large seed crops produced by Pacific dogwood. Following monitoring in Idaho, researchers found Pacific dogwood produced a significant amount of seed in only 1 year out of 5 [82]. Roper [105] reports that Pacific dogwood reproduces by seed only when growing under canopy cover.
It is likely that seed production is linked to plant maturity. Only larger trees produced seed in Idaho [82]. Roof [104] suggests that Pacific dogwood flowers first when approximately 6 years old. Others report that the minimum seed-bearing age of Pacific dogwood is 10 years [21].
Seed dispersal: The fleshy fruit surrounding Pacific dogwood seeds is likely attractive to bird and small mammal seed dispersers, but seed dispersal was not directly discussed in the literature cited as of this writing (2005).
Seed banking: Pacific dogwood likely banks some seed. In a seed production and seed bank study along 3rd and 5th order streams, Pacific dogwood coverage was recorded only in old-growth Douglas-fir sites on the 5th order stream, but 1 Pacific dogwood seed germinated in soil collected along the 3rd order stream [54].
Germination: Germination of Pacific dogwood seed under controlled conditions is high. When seed collected from Idaho populations was sent to horticulturalists, they reported a 63% germination rate [82]. In a review, Brinkman and Vankus [21] report that germination averaged 81% when tested on sand and wet paper.
Seedling establishment/growth: While shading seems important to seedling emergence, deep shade may not provide for establishment, growth, and reproduction of Pacific dogwood. Brush [23] reported that the highest number of seedlings grew in deep shade or on moist streambanks. Likewise, Roper [105] chiefly observed seedlings on sites with greater than 46% canopy cover. Successful seedling establishment in Idaho occurred on sites with 45%-60% canopy cover [105]. However more recently in Idaho, researchers could not find Pacific dogwood populations considered large enough for monitoring purposes in understory sites. Large populations of Pacific dogwood were associated with shrub-dominated areas. A single monitoring plot was located on a deep shade site, but plants had not recruited vegetatively or by seed in 5 years of monitoring [82].
Asexual regeneration:
Pacific dogwood readily sprouts following disturbance. In Idaho, sprouting has
been the only regeneration in several years, as no seedlings were located in monitoring
sites [82]. Two years following a clearcut operation in northwestern California,
Pacific dogwood averaged 19 sprouts per stump. Sprouts reached a maximum of 4.2 feet
(1.3 m) tall and clumps of sprouts were a maximum of 4.5 feet (1.4 m) in diameter [106].
Layering
was also observed in newly sprouting Pacific dogwood plants [105].
SITE CHARACTERISTICS:
Pacific dogwood's low frost tolerance, high flood tolerance, and moderate
shade tolerance make it common along stream banks and in low-elevation
coniferous, hardwood, and mixed coastal forests with temperate to
mesothermal climates [64,67,78,98]. Brush [23] considers Pacific dogwood
populations best developed in Douglas-fir forests of the Puget Sound Basin and
redwood forests of California. Typical habitat for Pacific dogwood includes
sites with moist but well-drained soils, on gentle slopes, predominantly
occurring below 5,000 feet (1,524 m) in elevation [23,98].
Climate: The climates in Pacific dogwood's range are mild and moist. Mediterranean-marine and temperate maritime weather patterns are typical [44,113,130]. Average annual precipitation ranges from a low of 12.2 inches (310 mm) in the Siskiyou Mountains of Oregon and California to a high of 100.4 inches (2,550 mm) in western Washington [17,133]. The majority of precipitation falls between late fall and early spring as rain and/or snow depending on the elevation of the site. During the growing season in parts of Washington, precipitation levels are less than 10% to 25% of the annual average [17,43]. Low winter temperatures range from 30.2°F (-1 °C) to 50 °F (10°C) reported for the Sierra Nevada and coasts of Oregon and California, respectively [16,107]. Maximum summer temperatures range from 60 to 72.7 °F (15.6-22.6 °C) reported for the coasts and Siskiyou Mountains of Oregon and California, respectively [107,133]. In Idaho, Pacific dogwood's restricted range is driven by climatic factors. The Lochsa-Selway region of Idaho receives high levels of winter and spring precipitation while summers can be hot and dry; annual precipitation in this area averages 34 inches (864 mm). Winter low temperatures average 29.5 °F (-1.4 °C) to 31.6 °F (-0.2 °C) and summer highs average 70 to 72°F (21-22.2 °C) [105].
Soils: The soils described in association with Pacific dogwood habitat are typically deep (often ≥ 6.6 feet (2 m)), moist, and well-drained [4,23,43,85,104]. Soil textures can range from clay to sand loam types [4,8,43]. Pacific dogwood soils often are high in humus content and have a low pH (5.5-6) [8]. The permeability of most Pacific dogwood soils is slow and the water holding capacity is high (59.1 to 98.4 inches (150-250 cm)) [4]. Moderate to high levels of calcium, magnesium, nitrates, potassium, and phosphorus are also typical of soils supporting Pacific dogwood [78].
While a majority of soils associated with Pacific dogwood are deep, in a study of early succession of the mud flows that followed the Mt. St. Helens volcanic eruption, Pacific dogwood was only found on nonorganic substrates with less than 9.8 inches (24.9 cm) of buried soil. Frequency of Pacific dogwood on this site was less than 1% [53].
Elevation:
California | Sierra Nevada Range: below 1,500 feet (547 m) [16] montane coniferous forests: below 6,500 feet (1,981 m) [91] |
California/Oregon | Coast Range: 492 to 1,969 feet (150-600 m) [113] Klamath Range: 1,969 to 5,249 feet (600-1600 m) [114] redwood forests: below 3,000 feet (914 m) [107] Siskiyou region: below 3,500 feet (1,067 m) [133] |
Idaho | Douglas-fir forests: 2,100 feet (366 m), no range was given [59] western redcedar forests: 2,500 to 1,421 feet (433-762 m) [101,105] |
Oregon | Cascade Range: 1,411 to 2,198 feet (430-670 m) [49] |
Washington | Mount Rainier: 1,804 to 3,543 feet (550-1080 m) [44] Wind River area: 1,099 to 2,362 feet (335-720 m) [17,43] |
In Idaho, Pacific dogwood was present in all stages of succession. Its prolific sprouting following the removal of above ground biomass is considered important in the recolonization of sites with pioneer conditions. Below are the average percent coverages of Pacific dogwood in order of advancing successional communities on western redcedar-western hemlock forest sites. In those communities for which a range of coverages is given, the range is representative of the average percent cover reported for multiple sites within the seral community [105].
Seral community | Immature shrub | Mature shrub | Mature shrub-young conifer | Shrub-Betula (birch) | Seral conifer | Climax conifer |
Pacific dogwood cover (%) | 8-19 | <1-5 | 31-48 | 9-21 | 16-88 | 3-7 |
When studies compared disturbed and undisturbed sites, Pacific dogwood was commonly present in both. Following a clearcut of 125-year-old Douglas-fir stands, Pacific dogwood was present the 3rd postlogging year [66]. Likewise, 5 years after Douglas-fir forests of northwestern California were logged Pacific dogwood coverage was 11%. In unlogged "virgin" forests, coverage of Pacific dogwood was 9% [51]. Pacific dogwood has even been considered an "invader" by some. In clearcut, old-growth Douglas-fir forests of the Cascade Range, Pacific dogwood was considered an invader because it was not found in the understory of adjacent areas. However, species frequencies were not measured prior to logging, making the absence of Pacific dogwood prior to logging an assumption [135].
In the Cascade Range of Oregon and Washington and in the Coast Range of Oregon, different-aged stands of Douglas-fir resulting from past fires were compared. The basal area of Pacific dogwood trees with greater than 2 inches (5 cm) d.b.h. was measured in young (< 80 years), mature (80-195 years), and old-growth (≥ 195 years) Douglas-fir forests. This study revealed no clear successional pattern for Pacific dogwood. The results are presented below [118]:
Cascade Range, WA | Cascade Range, OR | Coast Range, OR | |
Young | <0.1 m2/ha | 0.3 m2/ha | 0.2 m2/ha |
Mature | 0.2 m2/ha | 0.1 m2/ha | 0.1 m2/ha |
Old growth | <0.1 m2/ha | 0.1 m2/ha | 0.4 m2/ha |
Fire regimes: The fire regime for Pacific dogwood is dependent on the overstory community, site conditions, and historical disturbances. In the central and south Sierra Nevada, indigenous people historically burned areas to encourage new growth in Cornus spp. Fires were set in the fall and burning occurred at 1- to 2-year intervals [5]. Others also suggest that fires were common in the Sierra Nevada. Parsons and DeBenedetti [95] suggest that fires frequently burned in sequoia and mixed conifer forest types. The poor recruitment of giant sequoia is thought to be related to fire suppression efforts in these areas [95,129]. Fires were also frequent in the Siskiyou region of California and Oregon; fire severities however ranged widely [133].
In the Klamath Mountains of California, researchers investigated 75 plots in 3,880 acres (1,570 ha) to reconstruct the fire history. They estimated the average area burned was 865 acres (350 ha) with 16 fires between 1627 and 1992 that were greater than 1,236 acres (500 ha). Most fires were of low and moderate severities, although stand-replacing fires also occurred. Estimated average fire return intervals are shown below [124].
Presettlement (1626-1849) | 14.5 years |
Settlement (1850-1904) | 12.5 years |
Suppression (1905-1992) | 21.8 years |
In the Lochsa-Selway area of Idaho, Roper [105] reports that large fires burned in 1910, 1919, 1924, 1930, 1934 and smaller fires occurred in 1949-50 and 1967. Heavy moisture in the winter and spring months allows fuels to accumulate in this area, while hot, dry summers foster burning conditions. Climate regime is likely the primary driving factor of frequent fires in this area [105].
Not all areas where Pacific dogwood is common burned often. Riparian areas often burn less frequently and/or burn at lower severity than the surrounding slopes [1]. In coastal redwood forests of northern California, lightning caused fires were infrequent. In this high humidity region, the author estimates from age class and fire scar distributions that low-severity fires occurred at 250- to 500-year intervals on mesic sites, at 50-year intervals on xeric sites, and at 100- and 200-year intervals on intermediate sites [130].
The following table provides fire return intervals for plant communities and ecosystems where Pacific dogwood is important. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
Community or Ecosystem | Dominant Species | Fire Return Interval Range (years) |
silver fir-Douglas-fir | Abies amabilis-Pseudotsuga menziesii var. menziesii | > 200 |
grand fir | Abies grandis | 35-200 |
Engelmann spruce-subalpine fir | Picea engelmannii-Abies lasiocarpa | 35 to > 200 |
Jeffrey pine | Pinus jeffreyi | 5-30 |
western white pine* | Pinus monticola | 50-200 |
Pacific ponderosa pine* | Pinus ponderosa var. ponderosa | 1-47 [9] |
interior ponderosa pine* | Pinus ponderosa var. scopulorum | 2-30 [9,14,80] |
Rocky Mountain Douglas-fir* | Pseudotsuga menziesii var. glauca | 25-100 [9,10,12] |
coastal Douglas-fir* | Pseudotsuga menziesii var. menziesii | 40-240 [9,89,103] |
California mixed evergreen | Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii | < 35 |
California oakwoods | Quercus spp. | < 35 [9] |
coast live oak | Quercus agrifolia | 2-75 [48] |
canyon live oak | Quercus chrysolepis | <35 to 200 |
Oregon white oak | Quercus garryana | < 35 [9] |
California black oak | Quercus kelloggii | 5-30 [97] |
redwood | Sequoia sempervirens | 5-200 [9,41,122] |
western redcedar-western hemlock | Thuja plicata-Tsuga heterophylla | > 200 |
western hemlock-Sitka spruce | Tsuga heterophylla-Picea sitchensis | > 200 |
mountain hemlock* | Tsuga mertensiana | 35 to > 200 [9] |
In the western Cascade Mountains of Oregon 100- to 500-year-old Douglas-fir stands were clearcut then burned the following year to determine early successional development of these forests [36,37]. Prior to any treatments, Pacific dogwood coverage was 6.4% and 3.6% in vine maple-salal- and western sword fern-dominated communities, respectively. Five years after burning slash in these areas, Pacific dogwood coverage was 0.4% in vine maple-salal forest communities. In western sword fern communities that occupied stream banks and northern slopes, the coverage of Pacific dogwood was 1.4% in 5-year-old burned clearcuts [37].
Ingram [70] compared single- and multiple-slash burns in the Douglas-fir forests of the Columbia National Forest. Pacific dogwood density and percent composition were greater on sites that were burned the same year they were logged when compared to sites burned 1 year following logging and those sites burned twice after logging.
In the H.J. Andrews Experimental Forest of west-central Oregon, old growth stands of Douglas-fir (>400 years) were clearcut in June and slash burned in October. Pacific dogwood had 1% cover on the sites before being logged and burned; at the end of the next growing season Pacific dogwood coverage was either 0% or lower than the 1% cutoff used to warrant reporting. There were no control comparisons available for Pacific dogwood in this study [46].
In the western Cascade Mountains of Oregon, researchers studied secondary succession following clearcutting and broadcast burning of multiple sites within Douglas-fir forests. Pacific dogwood cover increased as secondary succession progressed. Results are presented below [116]:
Number of years since clearcut and broadcast burned | 2 | 5 | 10 | 15 | 20 | 30 | 40 | Undisturbed (450-year-old stand) |
Mean Pacific dogwood cover values (n = 3), |
0.04 | .013 | 0 | 0.04 | 0.52 | 0.17 | 1.69 | 0.85 |
Fire alone: Considerably fewer studies relate to the postfire response of Pacific dogwood to fire without logging as an additional major disturbance. After the large Oxbow fire that burned portions of western Oregon, Hooven [65] lists Pacific dogwood as one of many species that made up the postburn vegetation the 1st year following the fire. There was no information provided about fire severity or season of the burn.
In mixed conifer forests of the northern Sierra Nevada, California, 4 prescription fires burned on 2 sites where Pacific dogwood occurred. Fires were described as early spring-moderate consumption, late spring-high consumption, early fall-moderate consumption, and late fall-high consumption. Early spring prescribed fires coincided with bud break of most species, late spring burns occurred when plants were actively growing, early fall fires coincided with the end of the shrub growing season, and late fall burns corresponded with the leaf drop of most deciduous shrub species. Following late spring, early fall, and late fall fires, Pacific dogwood density increased. The early spring fire, however, had the opposite effect. The following table reports the effects of fire seasonality on Pacific dogwood; reported are the densities (number of plants/hectare) [74].
Pre-burn | 1st postburn growing season | 2nd postburn growing season | |
Early spring | 33 | 0 | 0 |
Late spring | 0 | 0 | 34 |
Early fall | 33 | -- | 67 |
Late fall | 0 | 42 | 125 |
The Research Project Summary Plant response to prescribed burning with varying season, weather, and fuel moisture in mixed-conifer forests of California provides information on prescribed fire and postfire response of many plant community species including Pacific dogwood.
FIRE MANAGEMENT CONSIDERATIONS:When setting prescription fires in giant sequoia groves, the California Parks and Recreation Department 1st raked around Pacific dogwood as they considered this species prone to cambium damage and wanted to reduce the "visual and environmental" impacts of the fire [57].
Domestic livestock: The quality of mature Pacific dogwood browse is considered fair to poor for domestic sheep and goats and is thought to be "worthless" browse for horses and cattle. New sprouts however are preferred by livestock [111]. Pacific dogwood was heavily grazed by domestic sheep in areas of the Columbia National Forest that had burned 4 to 8 years earlier [70].
Wildlife: Large mammals: In the summer, grazing by mule deer was greater than 50% (significantly greater (p<0.01)) for 3 sampling seasons following logging and burning [81]. Pacific dogwood sprouts resulting from a clearcut were utilized by deer. Sixty-two percent of sprouting clumps were browsed; 10% were considered heavily browsed [106]. Researchers found Pacific dogwood in 33% of the 69 elk stomachs collected from January through March near the Lochsa and Selway rivers of northern Idaho. However, Pacific dogwood browse made up just 3% of their total diet [127]. On a 40-year-old burn, black-tailed deer very rarely browsed Pacific dogwood even though food was scarce due to high deer populations (>100 deer/mi2) [61]. When browsing was monitored in enclosed Douglas-fir plantations burned 21-22 years ago, the high density black-tailed deer population (126 deer/mi2) did not feed on Cornus spp. during winter months [62].
Small mammals: Few studies have focused on small mammal use of Pacific dogwood. During the fall months in Plumas County, California, Pacific dogwood fruits were eaten and 1 end of the seeds was gnawed, likely by deer mice [71]. Gilbert and Allwine [47] report finding the red tree vole often with medium-sized Pacific dogwood and other berry producing shrubs. While this finding may be related to food preferences of this species, the authors caution that this finding may be the result of a small sample size.
Birds: Pacific dogwood fruits are attractive to many birds [11]. Band-tailed pigeons and pileated woodpeckers feed on Pacific dogwood fruits [11,87,98]
Palatability/nutritional value: Palatability of Pacific dogwood is associated with age of the plant tissue. Palatability is considered low for mature Cornus spp. due to bitter cell sap [111]. For slugs, the palatability of Pacific dogwood is considered average [27]. In a recently burned area however, Pacific dogwood is considered palatable [70]. Roper [105] claims that palatability of Pacific dogwood is highest for 2 or 3 years following fire.
Several nutrient and structural components of Pacific dogwood have been described. Pacific dogwood had the highest calcium levels of the 5 conifer, 4 shrub, and 2 broadleaf forest species analyzed in a giant sequoia forest community. Other nutrient levels reported were: 9% protein, 3.1%-2.7% fat, and 12.2%-13.6% fiber. Interestingly, calcium and fiber levels were slightly higher on logged and burned sites [81]. There are also high levels of aluminum in the leaves and branchlets of Pacific dogwood [102]. The average lignin and nitrogen content of leaves collected in Oregon was 6.2% and 0.87%, respectively [132].
Cover value:
Few studies have investigated the value of Pacific dogwood as habitat or
cover. It is probable though that this species provides cover and habitat to some
birds and small mammals. Pacific dogwood can grow as a large shrub or tree and
likely these different forms provide cover or habitat for different animal species. Sampson
[111] considers Cornus spp. important shade providers to larger ungulate species.
Wilson's warbler is attracted to meadows surrounded by Cornus spp. [119].
VALUE FOR REHABILITATION OF DISTURBED SITES:
Pacific dogwood is easily grown from seed and various seed treatments have been
described to artificially overcome dormancy [35,56,84]. Seed collected in the fall
can be sown directly into mineral soil to attain the long cool stratification required to
overcome seed dormancy. Seed collected in the summer can be dried and refrigerated until
fall [56]. Others have soaked seed in concentrated sulfuric acid to overcome seed
dormancy [35,84].
The use of Pacific dogwood in revegetation efforts has appealed to many. Evaluations of the revegetation potential of Pacific dogwood have been undertaken as well [34,99].
There are other aspects of Pacific dogwood biology that may affect its use in
rehabilitation or revegetation efforts. Pacific dogwood leaves decay rapidly [55,132].
Decay rates for Pacific dogwood were faster than any of the other 6 species tested. Tinnin
and Kirkpatrick [126] assessed the growth suppression potential of Pacific dogwood. In
a greenhouse study, they found radicle growth to be 38% of normal when cucumber
seed was grown on sponges soaked with water leachates of Pacific dogwood leaf
litter. The use of cucumber seed in this study makes it difficult at best to
make any inferences regarding Pacific dogwood's ability to suppress any native
or naturally occurring plant neighbors.
OTHER USES:
Showy flowers and brilliant fall colors make Pacific dogwood a valuable
ornamental species [104,112].
Pacific dogwood bark was used by Nlaka `pamux, indigenous people of the Pacific Northwest Coast, to make brown dye. Bark has also been prepared and used as a blood purifier, lung strengthener, and stomach treatment [98]. Arno [11] suggests that historically the bark of Cornus spp. was used to cure malaria and when boiled had laxative properties.
Wood Products: The wood of Pacific dogwood has several uses. This hardwood species has been used to make bows, arrows, thread spindles, cabinets, piano keys, mallet handles, golf club heads and other tools [2,11,23,67,98]. Young shoots of Cornus spp. were used by indigenous people of central and south Sierra Nevada for basket making [5]. The collection of Pacific dogwood is currently prohibited in British Columbia [98].
The wood of Pacific dogwood is hard, heavy, has a whitish color, a fine grain, and wears
smoothly [67,93]. For more about wood properties see [2,23,93,96] and for treatment of the
wood see [2,93].
OTHER MANAGEMENT CONSIDERATIONS:
Managers and researchers have found many uses for Pacific dogwood. Klinka and
others [77] found Pacific dogwood to be one of many species that indicates cool
mesothermal climates, nitrogen-rich soils, and soils with a moisture deficit <
3.5 months of the year. Minore [88] found Pacific dogwood was a valuable indicator of
summer soil temperatures and moisture stress in the southern Umpqua Basin of Oregon. Pacific
dogwood has also been used as a moisture index indicator species in mixed evergreen stands
of the Sierra Nevada [92]. Dawson and Greco [33] suggested protecting Pacific dogwood
in vegetation pockets while prescribe burning areas of Sequoia National Park to retain
aesthetics for visitors.
1. Agee, James K. 1994. Fire and weather disturbances in terrestrial ecosystems of the eastern Cascades. Gen. Tech. Rep. PNW-GTR-320. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 52 p. (Everett, Richard L., assessment team leader; Eastside forest ecosystem health assessment; Hessburg, Paul F., science team leader and tech. ed., Volume III: assessment). [23656]
2. Alden, Harry A. 1995. Hardwoods of North America. Gen. Tech. Rep. FPL-GTR-83. Madison, WI: U.S. Department of Agriculture, Forest Service, Forest Products Laboratory. 136 p. Available: http://www.fpl.fs.usda.gov/documnts/fplgtr/fplgtr83.pdf [2004, January 6]. [46270]
3. Anderson, Howard George. 1967. The phytosociology of some vine maple communities in the Mary's Peak watershed. Corvallis, OR: Oregon State University. 118 p. Thesis. [9877]
4. Anderson, John Melvin. 1982. Effects of prescribed burning on shrub seed stored in the duff and soil of a Sierra Nevada mixed-conifer forest. Berkeley, CA: University of California. 39 p. Thesis. [28337]
5. Anderson, M. Kat. 1996. Tending the wilderness. Restoration & Management Notes. 14(2): 154-166. [35819]
6. Anderson, R. Scott. 1990. Modern pollen rain within and adjacent to two giant sequoia (Sequoiadendron giganteum) groves, Yosemite and Sequoia National Parks, California. Canadian Journal of Forest Research. 20: 1289-1305. [15166]
7. Anderson, R. Scott. 1994. Paleohistory of a giant sequoia grove: the record from Log Meadow, Sequoia National Park. In: Aune, Philip S., technical coordinator. Proceedings of the symposium on giant sequoias: their place in the ecosystem and society; 1992 June 23-25; Visalia, CA. Gen. Tech. Rep. PSW-GTR-151. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 49-55. [24749]
8. Andrews, Susyn. 1991. Cornus nuttallii in cultivation. The Kew Magazine. 8(2): 71-78. [49484]
9. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]
10. Arno, Stephen F.; Gruell, George E. 1983. Fire history at the forest-grassland ecotone in southwestern Montana. Journal of Range Management. 36(3): 332-336. [342]
11. Arno, Stephen F.; Hammerly, Ramona P. 1977. Northwest trees. Seattle, WA: The Mountaineers. 222 p. [4208]
12. Arno, Stephen F.; Scott, Joe H.; Hartwell, Michael G. 1995. Age-class structure of old growth ponderosa pine/Douglas-fir stands and its relationship to fire history. Res. Pap. INT-RP-481. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 25 p. [25928]
13. Atzet, Tom; Wheeler, David; Smith, Brad; [and others]. 1985. The tanoak series of the Siskiyou region of southwest Oregon (Part 2). Forestry Intensified Research. 6(4): 7-10. [8594]
14. Baisan, Christopher H.; Swetnam, Thomas W. 1990. Fire history on a desert mountain range: Rincon Mountain Wilderness, Arizona, U.S.A. Canadian Journal of Forest Research. 20: 1559-1569. [14986]
15. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
16. Bias, Michael A.; Gutierrez, R. J. 1992. Habitat associations of California spotted owls in the central Sierra Nevada. Journal of Wildlife Management. 56(3): 584-595. [19279]
17. Bishaw, Badege; DeBell, Dean S.; Harrington, Constance A. 2003. Patterns of survival, damage, and growth for western white pine in a 16-year-old spacing trial in western Washington. Western Journal of Applied Forestry. 18(1): 35-43. [44996]
18. Biswell, H. H. 1959. Prescribed burning and other methods of deer range improvement in ponderosa pine in California. In: Proceedings, Society of American Foresters; 1959; San Francisco, CA. Bethesda, MD: Society of American Foresters: 102-105. [5269]
19. Biswell, H. H.; Schultz, A. M.; Launchbaugh, J. L. 1955. Brush control in ponderosa pine. California Agriculture. 9(1): 3, 14. [16358]
20. Boroski, Brian B.; Barrett, Reginald H.; Timossi, Irene C.; Kie, John G. 1996. Modelling habitat suitability for black-tailed deer (Odocoileus hemionus columbianus) in heterogeneous landscapes. Forest Ecology and Management. 88: 157-165. [27307]
21. Brinkman, Kenneth A.; Vankus, Victor. 2004. Cornus L. dogwood, [Online]. In: Bonner, Franklin T., tech. coord. Woody plant seed manual. Washington, DC: U.S. Department of Agriculture, National Tree Seed Laboratory (Producer). Available: http://ntsl.fs.usda.gov/wpsm/Cornus.pdf [2005, January 27]. [51535]
22. Brown, Daniel A.; Windham, Mark T.; Trigiano, Robert N. 1996. Resistance to dogwood anthracnose among Cornus species. Journal of Arboriculture. 22(2): 83-86. [48969]
23. Brush, Warren. 1948. Pacific dogwood: Cornus nuttalli, Audubon. American Forests. 54(5): 216-217. [49487]
24. Byther, Ralph S.; Davidson, Roy M., Jr. 1994. Dogwood anthracnose. Extension Bulletin 0972 [revised]. Pullman, WA: Washington State University, College of Agriculture and Home Economics, Cooperative Extension. 2 p. [49496]
25. Caetano-Anolles, Gustavo; Trigiano, Robert N.; Windham, Mark T. 2001. Patterns of evolution in Discula fungi and the origin of dogwood anthracnose in North America, studied using arbitrarily amplified and ribosomal DNA. Current Genetics. 39(5-6): 346-354. [48966]
26. Carey, Andrew B.; Kershner, Janet; Biswell, Brian; Dominguez de Toledo, Laura. 1999. Ecological scale and forest development: squirrels, dietary fungi, and vascular plants in managed and unmanaged forests. Wildlife Monographs. 142: 1-71. [30476]
27. Cates, Rex G.; Orians, Gordon H. 1975. Successional status and the palatability of plants to generalized herbivores. Ecology. 56: 410-418. [15989]
28. Chambers, Carol L.; Carrigan, Tara; Sabin, Thomas E.; [and others]. 1997. Use of artificially created Douglas-fir snags by cavity-nesting birds. Western Journal of Applied Forestry. 12(3): 93-97. [27530]
29. Daubenmire, R. 1969. Ecologic plant geography of the Pacific Northwest. Madrono. 20: 111-128. [740]
30. Daughtrey, M. L.; Hibben, C. R. 1994. Dogwood anthracnose: a new disease threatens two native Cornus species. Annual Review of Phytopathology. 32: 61-73. [49482]
31. Daughtrey, Margery L.; Hibben, Craig R.; Britton, Kelly O.; Windham, Mark T.; Redlin, Scott C. 1996. Dogwood anthracnose: Understanding a disease new to North America. Plant Disease. 80(4): 349-358. [48967]
32. Davidson, Roy M., Jr.; Byther, Ralph S. 1981. Dogwood anthracnose. Extension Bulletin EB 0972. Pullman, WA: Washington State University, College of Agriculture, Cooperative Extension. 2 p. [49459]
33. Dawson, Kerry J.; Greco, Steven E. 1991. Prescribed fire and visual resources in Sequoia National Park. In: Nodvin, Stephen C.; Waldrop, Thomas A., eds. Fire and the environment: ecological and cultural perspectives: Proceedings of an international symposium; 1990 March 20-24; Knoxville, TN. Gen. Tech. Rep. SE-69. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 192-201. [16650]
34. DeBolt, Ann; Spurrier, Carol S. 2004. Seeds of success and the Millennium Seed Bank project. In: Hild, Ann L.; Shaw, Nancy L.; Meyer, Susan E.; Booth, D. Terrance; McArthur, E. Durant, compilers. Seed and soil dynamics in shrubland ecosystems: proceedings; 2002 August 12-16; Laramie, WY. Proceedings RMRS-P-31. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 100-108. [49104]
35. Dummer, Peter. 1969. Problems in raising ornamental stock from seed. Proceedings, Plant Propagation Society. 19: 213-215. [49483]
36. Dyrness, C. T. 1965. The effect of logging and slash burning on understory vegetation in the H. J. Andrews Experimental Forest. Res. Note PNW-31. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 13 p. [4939]
37. Dyrness, C. T. 1973. Early stages of plant succession following logging and burning in the western Cascades of Oregon. Ecology. 54(1): 57-69. [7345]
38. Edson, John L.; Wenny, David L.; Leege-Brusven, Annette. 1994. Micropropagation of Pacific dogwood. HortScience. 29(1): 1355-1356. [48960]
39. Evenden, Angela G. 1990. Sensitive plants and natural areas. In: Evenden, Angela G., compiler. Proceedings--Northern Region biodiveristy workshop; 1990 September 11-13; Missoula, MT. Missoula, MT: U.S. Department of Agriculture, Forest Service, Northern Region: 36-39. [23594]
40. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
41. Finney, Mark A.; Martin, Robert E. 1989. Fire history in a Sequoia sempervirens forest at Salt Point State Park, California. Canadian Journal of Forest Research. 19: 1451-1457. [9845]
42. Flora of North America Association. 2000. Flora of North America north of Mexico. Volume 2: Pteridophytes and gymnosperms, [Online]. Flora of North America Association (Producer). Available: http://hua.huh.harvard.edu/FNA/ [2004, October 27]. [36990]
43. Franklin, Jerry F.; DeBell, Dean S. 1988. Thirty-six years of tree population change in an old-growth Pseudotsuga-Tsuga forest. Canadian Journal of Forest Research. 18: 633-639. [8769]
44. Franklin, Jerry F.; Moir, William H.; Hemstrom, Miles A.; [and others]. 1988. The forest communities of Mount Rainier National Park. Scientific Monograph Series No 19. Washington, DC: U.S. Department of the Interior, National Park Service. 194 p. [12393]
45. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
46. Gashwiler, Jay S. 1959. Small mammal study in west-central Oregon. Journal of Mammalogy. 40(1): 128-139. [14005]
47. Gilbert, Frederick F.; Allwine, Rochelle. 1991. Small mammal communities in the Oregon Cascade Range. In: Ruggiero, Leonard F.; Aubry, Keith B.; Carey, Andrew B.; Huff, Mark H., technical coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 257-267. [17317]
48. Greenlee, Jason M.; Langenheim, Jean H. 1990. Historic fire regimes and their relation to vegetation patterns in the Monterey Bay area of California. The American Midland Naturalist. 124(2): 239-253. [15144]
49. Grier, Charles C.; Logan, Robert S. 1977. Old-growth Pseudotsuga menziesii communties of a western Oregon watershed: biomass distribution and production budgets. Ecological Monographs. 47: 373-400. [8762]
50. Habeck, James R. 1961. The original vegetation of the mid-Willamette Valley, Oregon. Northwest Science. 35: 65-77. [11419]
51. Hagar, Donald C. 1960. The interrelationships of logging, birds, and timber regeneration in the Douglas-fir region of northwestern California. Ecology. 41(1): 116-125. [34500]
52. Halpern, C. B. 1989. Early successional patterns of forest species: interactions of life history traits and disturbance. Ecology. 70(3): 704-720. [6829]
53. Halpern, Charles B.; Harmon, Mark E. 1983. Early plant succession on the Muddy River mudflow, Mount St. Helens, Washington. The American Midland Naturalist. 110(1): 97-106. [8870]
54. Harmon, Janice M.; Franklin, Jerry F. 1995. Seed rain and seed bank of third- and fifth-order streams on the western slope of the Cascade Range. Res. Pap. PNW-RP-480. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 27 p. [26575]
55. Harmon, Mark E.; Baker, Gail A.; Spycher, Gody; Greene, Sarah E. 1990. Leaf-litter decomposition in the Picea/Tsuga forests of Olympic National Park, Washington, U. S. A. Forest Ecology and Management. 31: 55-66. [10318]
56. Harrington, Constance A.; McGrath, James M.; Kraft, Joseph M. 1999. Propagating native species: experience at the Wind River Nursery. Western Journal of Applied Forestry. 14(2): 61-64. [30058]
57. Harrison, Wayne. 1986. Management of giant sequoia at Calaveras Big Trees State Park. In: Weatherspoon, C. Phillip; Iwamoto, Y. Robert; Piirto, Douglas D., technical coordinators. Proceedings of the workshop on management of giant sequoia; 1985 May 24-25; Reedley, CA. Gen. Tech. Rep. PSW-95. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 40-42. [9811]
58. Hawk, G. M.; Zobel, D. B. 1974. Forest succession on alluvial landforms of the McKenzie River Valley, Oregon. Northwest Science. 48(4): 245-265. [9686]
59. Hickey, William O. 1971. Response of Ceanothus sanguineus to cutting and burning at various stages of phenology. Moscow, ID: University of Idaho. 39 p. Thesis. [9777]
60. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
61. Hines, William W. 1973. Black-tailed deer populations and Douglas-fir reforestation in the Tillamook Burn, Oregon. Game Research Report Number 3. Federal Aid to Wildlife Restoration, Project W-51-R: Final Report. Corvallis, OR: Oregon State Game Commission. 59 p. [8431]
62. Hines, William W.; Land, Charles E. 1974. Black-tailed deer and Douglas-fir regeneration in the Coast Range of Oregon. In: Black, Hugh C., ed. Wildlife and forest management in the Pacific Northwest: Proceedings of a symposium; 1973 September 11-12; Corvallis, OR. Corvallis, OR: Oregon State University, School of Forestry, Forest Research Laboratory: 121-132. [7999]
63. Hitchcock, C. Leo; Cronquist, Arthur. 1961. Vascular plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaceae. Seattle, WA: University of Washington Press. 614 p. [1167]
64. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. [1168]
65. Hooven, Edward F. 1969. The influence of forest succession on populations of small animals in western Oregon. In: Black, Hugh C., ed. Wildlife and reforestation in the Pacific Northwest: Proceedings of a symposium; 1968 September 12-13; Corvallis, OR. Corvallis, OR: Oregon State University, School of Forestry: 30-34. [7943]
66. Hooven, Edward F. 1973. Response of the Oregon creeping vole to the clearcutting of a Douglas-fir forest. Northwest Science. 47(4): 256-264. [8521]
67. Hosie, R. C. 1969. Native trees of Canada. 7th ed. Ottawa, ON: Canadian Forestry Service, Department of Fisheries and Forestry. 380 p. [3375]
68. Idaho Fish and Game. 2005. Idaho's special status vascular plants, [Online]. Idaho Conservation Data Center (Producer). Available: http://fishandgame.idaho.gov/tech/DCD/plants/home.cfm [2005, January 24]. [51537]
69. Idaho Native Plant Society. 2004. The Idaho Native Plant Society rare plant list: State rare species list, [Online]. In: Results of the 20th annual Idaho rare plant conference. Idaho Native Plant Society (Producer). Available: http://www.idahonativeplants.org/rpc/RarePlantList.aspx [2005, January 24]. [51534]
70. Ingram, Douglas C. 1931. Vegetative changes and grazing use on Douglas-fir cut-over land. Journal of Agricultural Research. 43(5): 387-417. [8877]
71. Jameson, E. W., Jr. 1952. Food of deer mice, Peromyscus maniculatus and P. boylei, in the northern Sierra Nevada, California. Journal of Mammalogy. 33(1): 50-60. [21605]
72. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with the Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. [36715]
73. Kauffman, J. B.; Martin, R. E. 1990. Sprouting shrub response to different seasons and fuel consumption levels of prescribed fire in Sierra Nevada mixed conifer ecosystems. Forest Science. 36(3): 748-764. [13063]
74. Kauffman, John Boone. 1986. The ecological response of the shrub component to prescribed burning in mixed conifer ecosystems. Berkeley, CA: University of California, Berkeley. 235 p. Dissertation. [19559]
75. Kilgore, Bruce M. 1971. Response of breeding bird populations to habitat changes in a giant sequoia forest. The American Midland Naturalist. 85(1): 135-152. [7281]
76. King, David A. 1991. Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon. Tree Physiology. 9(3): 369-381. [48473]
77. Klinka, K.; Krajina, V. J.; Ceska, A.; Scagel, A. M. 1989. Indicator plants of coastal British Columbia. Vancouver, BC: University of British Columbia Press. 288 p. [10703]
78. Krajina, V. J.; Klinka, K.; Worrall, J. 1982. Distribution and ecological characteristics of trees and shrubs of British Columbia. Vancouver, BC: University of British Columbia, Department of Botany and Faculty of Forestry. 131 p. [6728]
79. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]
80. Laven, R. D.; Omi, P. N.; Wyant, J. G.; Pinkerton, A. S. 1980. Interpretation of fire scar data from a ponderosa pine ecosystem in the central Rocky Mountains, Colorado. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 46-49. [7183]
81. Lawrence, George; Biswell, Harold. 1972. Effect of forest manipulation on deer habitat in giant sequoia. Journal of Wildlife Management. 36(2): 595-605. [41671]
82. Lichthardt, Juanita. 1996. Population monitoring of disjunct Pacific dogwood (Cornus nuttallii) on the Clearwater National Forest: five year summary, [Online]. In: Botany publications--Idaho Conservation Data Center. Boise, ID: Idaho Fish and Game (Producer). Available: http://fishandgame.id.gov;tech/CDC/cdc_pdf/CORNUS95.PDF [2005, January 24]. [51538]
83. Lynch, John A. 1981. The outriders of spring: Dogwood. American Forests. 87(4): 22-27. [14095]
84. Macdonald, Bruce. 1990. Ornamental native plants of British Columbia: their selection, propagation, and introduction. Proceedings, International Plant Propagators' Society. 39: 243-249. [49460]
85. Marston, Ted. 1992. Cornus spp.: dogwood. Arbor Age. 12(9): 30-31. [20257]
86. McDonald, Philip M. 1980. Pacific ponderosa pine - Douglas-fir. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 120. [50055]
87. Michael, Charles W. 1928. The pileated woodpecker feeds on berries. The Condor. 30(2): 157. [8075]
88. Minore, Don. 1972. A classification of forest environments in the South Umpqua basin. Res. Pap. PNW-129. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 28 p. [1660]
89. Morrison, Peter H.; Swanson, Frederick J. 1990. Fire history and pattern in a Cascade Range landscape. Gen. Tech. Rep. PNW-GTR-254. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 77 p. [13074]
90. Moseley, Robert K. 1992. Rare plant conservation in Idaho. Journal of the Idaho Academy of Science. 28(2): 82-93. [22293]
91. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]
92. Myatt, Rodney G. 1980. Canyon live oak vegetation in the Sierra Nevada. In: Plumb, Timothy R., technical coordinator. Proceedings of the symposium on the ecology, management and utilization of California oaks; 1979 June 26-28; Claremont, CA. Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 86-91. [7019]
93. Overholser, J. L. 1977. Oregon hardwood timber. Corvallis, OR: Oregon State University, Forest Research Laboratory. 43 p. [16165]
94. Parsons, Dave. 1989. [Letter to Richard G. Krebill]. August 14. 1 leaf.. On file at: U.S. Department of Agriculture, Forest Service, Intermountain Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. [11440]
95. Parsons, David J.; DeBenedetti, Steven H. 1979. Impact of fire suppression an a mixed-conifer forest. Forest Ecology and Management. 2: 21-33. [7618]
96. Paul, Benson H. 1962. Choose the right wood: Properties and uses of some western hardwoods. Woodworking Digest. 64(3): 47-49. [49497]
97. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; [and others]. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
98. Pojar, Jim; MacKinnon, Andy, eds. 1994. Plants of the Pacific Northwest coast: Washington, Oregon, British Columbia and Alaska. Redmond, WA: Lone Pine Publishing. 526 p. [25159]
99. Radtke, Klaus. 1978. Wildland plantings & urban forestry: Native and exotic 1911-1977. Los Angeles, CA: County of Los Angeles Department of Forester and Fire Warden, Forestry Division. 134 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Forestry Research, Chaparral R & D Program. [20562]
100. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
101. Reel, Susan; Schassberger, Lisa; Ruediger, William. 1989. Caring for our natural community: Region 1 - Threatened, endangered, and sensitive species program. Missoula, MT: U.S. Department of Agriculture, Forest Service, Northern Region, Wildlife and Fisheries. 309 p. [51533]
102. Richardson, Andrew D.; Denny, Ellen G.; Forbush, Jocelyn A.; Siccama, Thomas G.; Hunter, Kimberly S. 2001. Differential aluminum and calcium concentrations in the tissues of ten Cornus species. Journal of the Torrey Botanical Society. 128(2): 120-127. [48987]
103. Ripple, William J. 1994. Historic spatial patterns of old forests in western Oregon. Journal of Forestry. 92(11): 45-49. [33881]
104. Roof, J. 1951. Growing California's five dogwoods. Journal of the California Horticulture Society. 12(3): 50-58, 101-106. [9002]
105. Roper, Laren Alden. 1970. Some aspects of the synecology of Cornus nuttallii in northern Idaho. Moscow, ID: University of Idaho. 81 p. Thesis. [51548]
106. Roy, D. F. 1955. Hardwood sprout measurements in northwestern California. PSW-95. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 6 p. [8999]
107. Roy, Douglas F. 1980. Redwood. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 109-110. [50047]
108. Rundel, Philip W. 1987. Origins and adaptations of California hardwoods. In: Plumb, Timothy R.; Pillsbury, Norman H., technical coordinators. Proceedings of the symposium on multiple-use management of California's hardwood resources; 1986 November 12-14; San Luis Obispo, CA. Gen. Tech. Rep. PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 11-17. [5355]
109. Rundel, Philip Wilson. 1969. The distribution and ecology of the giant sequoia ecosystem in the Sierra Nevada, California. Durham, NC: Duke University. 205 p. Dissertation. [37436]
110. Russel, D. W. 1974. The life history of vine maple on the H. J. Andrews Experimental Forest. Corvallis, OR: Oregon State University. 167 p. Thesis. [4974]
111. Sampson, Arthur W.; Jespersen, Beryl S. 1963. California range brushlands and browse plants. Berkeley, CA: University of California, Division of Agricultural Sciences, California Agricultural Experiment Station, Extension Service. 162 p. [3240]
112. Santamour, Frank S., Jr.; McArdle, Alice Jacot. 1985. Cultivar checklists of the large-bracted dogwoods: Cornus florida, C. kousa, and C. nuttallii. Journal of Arboriculture. 11(1): 29-36. [48968]
113. Sawyer, John O., Jr. 1980. Douglas-fir - tanoak - Pacific madrone. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 111-112. [50048]
114. Sawyer, John O.; Thornburgh, Dale A. 1977. Montane and subalpine vegetation of the Klamath Mountains. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 699-732. [685]
115. Sawyer, John O.; Thornburgh, Dale A.; Griffin, James R. 1977. Mixed evergreen forest. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 359-381. [7218]
116. Schoonmaker, Peter; McKee, Arthur. 1988. Species composition and diversity during secondary succession of coniferous forests in the western Cascade Mountains of Oregon. Forest Science. 34(4): 960-979. [6214]
117. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
118. Spies, Thomas A. 1991. Plant species diversity and occurrence in young, mature, and old-growth Douglas-fir stands in western Oregon and Washington. In: Ruggiero, Leonard F.; Aubry, Keith B.; Carey, Andrew B.; Huff, Mark H., technical coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 111-121. [17309]
119. Stebbins, C. A.; Stebbins, R. C. 1954. Birds of Yosemite National Park. Yosemite Nature Notes. 33(8): 74-152. [22717]
120. Stein, William I. 1980. Oregon white oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 110-111. [9857]
121. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [20090]
122. Stuart, John D. 1987. Fire history of an old-growth forest of Sequoia sempervirens (Taxodiaceae) forest in Humboldt Redwoods State Park, California. Madrono. 34(2): 128-141. [7277]
123. Tappeiner, John C. 1980. Sierra Nevada mixed conifer. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 118-119. [50054]
124. Taylor, Alan H.; Skinner, Carl N. 1998. Fire history and landscape dynamics in a late-successional reserve, Klamath Mountains, California, USA. Forest Ecology and Management. 111(2-3): 285-301. [30321]
125. Thilenius, John F. 1968. The Quercus garryana forests of the Willamette Valley, Oregon. Ecology. 49(6): 1124-1133. [8765]
126. Tinnin, Robert O.; Kirkpatrick, Lee Ann. 1985. The allelopathic influence of broadleaf trees and shrubs on seedlings of Douglas-fir. Forest Science. 31(4): 945-952. [9692]
127. Trout, Lester C.; Leege, Thomas A. 1971. Are the northern Idaho elk herds doomed? Idaho Wildlife Review. Nov-Dec: 3-6. [16731]
128. U.S. Department of Agriculture, National Resource Conservation Service. 2005. PLANTS database (2004), [Online]. Available: https://plants.usda.gov /. [34262]
129. Vankat, John L.; Major, Jack. 1978. Vegetation changes in Sequoia National Park, California. Journal of Biogeography. 5: 377-402. [17353]
130. Veirs, Stephen D., Jr. 1982. Coast redwood forest: stand dynamics, successional status, and the role of fire. In: Means, Joseph E., ed. Forest succession and stand development research in the Northwest: Proceedings of the symposium; 1981 March 26; Corvallis, OR. Corvallis, OR: Oregon State University, Forest Research Laboratory: 119-141. [4778]
131. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; [and others]. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]
132. White, G. J.; Baker, G. A.; Harmon, M. E.; Wiersma, G. B.; Bruns, D. A. 1990. Use of forest ecosystem process measurements in an integrated environmental monitoring program in the Wind River Range, Wyoming. In: Schmidt, Wyman C.; McDonald, Kathy J., compilers. Proceedings--symposium on whitebark pine ecosystems: ecology and management of a high-mountain resource; 1989 March 29-31; Bozeman, MT. Gen. Tech. Rep. INT-270. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 214-222. [11689]
133. Whittaker, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs. 30(3): 279-338. [6836]
134. Whittaker, R. H. 1961. Vegetation history of the Pacific Coast states and the "central" significance of the Klamath region. Madrono. 16: 5-23. [7467]
135. Yerkes, Vern P. 1960. Occurrence of shrubs and herbaceous vegetation after clear cutting old-growth Douglas-fir. Res. Pap. PNW-34. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 12 p. [8937]