SPECIES: Cornus nuttallii

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Cornus nuttallii: INTRODUCTORY

INTRODUCTORY

SPECIES: Cornus nuttallii

 

1995 Saint Mary's College of California

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: http://www.fs.fed.us/database/feis/ [].

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..."


DISTRIBUTION AND OCCURRENCE

SPECIES: Cornus nuttallii
GENERAL DISTRIBUTION:
Pacific dogwood occupies a discontinuous range. It occurs in the coastal regions west of the Cascade Mountains from southern British Columbia to southern California [39,64,91]. Only occasionally are Pacific dogwood populations found in the mountains of San Diego and Los Angeles counties of California [91]. There is a disjunct population of Pacific dogwood in northern Idaho along the lower Lochsa and Selway rivers [8,38,39,82]. The Idaho population is considered threatened [38,68,69,90].

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

CANADA
BC

BLM PHYSIOGRAPHIC REGIONS [15]:
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
8 Northern Rocky Mountains

KUCHLER [79] PLANT ASSOCIATIONS:
K001 Spruce-cedar-hemlock forest
K002 Cedar-hemlock-Douglas-fir forest
K003 Silver fir-Douglas-fir forest
K004 Fir-hemlock forest
K005 Mixed conifer forest
K006 Redwood forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K025 Alder-ash forest
K026 Oregon oakwoods
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods

SAF COVER TYPES [40]:
205 Mountain hemlock
206 Engelmann spruce-subalpine fir
210 Interior Douglas-fir
211 White fir
213 Grand fir
215 Western white pine
221 Red alder
222 Black cottonwood-willow
223 Sitka spruce
224 Western hemlock
225 Western hemlock-Sitka spruce
226 Coastal true fir-hemlock
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
231 Port-Orford-cedar
232 Redwood
233 Oregon white oak
234 Douglas-fir-tanoak-Pacific madrone
237 Interior ponderosa pine
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
249 Canyon live oak
255 California coast live oak

SRM (RANGELAND) COVER TYPES [117]:
109 Ponderosa pine shrubland
202 Coast live oak woodland
203 Riparian woodland

HABITAT TYPES AND PLANT COMMUNITIES:
British Columbia: Pacific dogwood is common in Douglas-fir (Pseudotsuga menziesii) and western hemlock (Tsuga heterophylla) forests of southwestern British Columbia [78].

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).


BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Cornus nuttallii

 

1998 Charles Webber, California Academy of Sciences

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 30.2F (-1 C) to 50 F (10C) 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 72F (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]

SUCCESSIONAL STATUS:
Pacific dogwood can tolerate early-, mid-, and late-seral conditions. In a study of herbivore successional preferences, researchers considered the presence of Pacific dogwood an indication of early-seral perennial vegetation based on reviewed literature and peak population occurrences [27]. However, in a study of water-formed terraces, floodplains, and glacial outwash plains along the McKenzie River Valley in Oregon, Pacific dogwood was present only in what the author considered late seral (100- to 200-year-old) and climax (200- to 500-year-old) communities [58]. Whittaker [134] considers Pacific dogwood part of the climax mixed evergreen vegetation in the Klamath region.

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

SEASONAL DEVELOPMENT:
Pacific dogwood commonly flowers twice in a growing season. This phenomenon may be related to late-summer water stress [104]. Flowers appear 1st in the spring (April, May, or June) [11,67]. In late summer or early fall, flowering often occurs again [23,67,91,98]. Fruits are often ripe by September or October [11,23,84]. In Idaho, Roper [105] reported that following exceptionally cold winter temperatures (minimum recorded was -12 F (-24 C)) May flowering was nearly eliminated.


FIRE ECOLOGY

SPECIES: Cornus nuttallii
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Pacific dogwood sprouts following fire [105]. Root crown sprouting and/or epicormic branching can occur following fire [1,19].

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. For further information, see the FEIS review of the dominant species listed below.

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]
*fire return interval varies widely; trends in variation are noted in the species review

POSTFIRE REGENERATION STRATEGY [121]:
Tree with adventitious bud/root crown
Crown residual colonizer (on-site, initial community)


FIRE EFFECTS

SPECIES: Cornus nuttallii
IMMEDIATE FIRE EFFECT ON PLANT:
The immediate effect of fire on Pacific dogwood varies with fire severity. Low-consumption, early-spring fires, in mature mixed conifer ecosystems of the northern Sierra Nevada, produced temperatures hot enough to kill Pacific dogwood foliage but not hot enough to kill buds protected by bark [73].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Buds that survive a low-severity fire are released and able to regenerate [1,73].

PLANT RESPONSE TO FIRE:
After being burned, Pacific dogwood typically sprouts from the root crown [105]. Parsons in a personal communication [94] predicts Pacific dogwood will sprout following fire "regardless of burn prescription." When 300- to 500-year-old Douglas-fir stands were logged and burned, Pacific dogwood was classified as a residual colonizer by the author [52].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Fire and logging: Much of the research regarding the postfire recovery of Pacific dogwood involves the study of areas that were logged before being burned. Likely, fires in logged areas are different from those that burn standing forests. Following the burning of slash piles of a clearcut Douglas-fir forest type in the central Cascade Range, Pacific dogwood initially decreased in constancy and cover [52].

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),
in year 10 (n = 2)

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:
Fire management considerations often involve more than just the postfire community or species response. Following an early spring fire in giant sequoia forests of Tulare County, California, Lawrence and Biswell [81] found that the utilization of Pacific dogwood by mule deer was significantly greater (p<0.01) on logged and burned sites. The authors note that the "resulting crown sprouts were browsed so heavily that the survival of such trees seemed doubtful."

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].


MANAGEMENT CONSIDERATIONS

SPECIES: Cornus nuttallii
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Utilization of Pacific dogwood by large mammals and livestock is related to time since last disturbance, as new sprouts are browsed more heavily than mature vegetation.

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.


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