Calocedrus decurrens



INTRODUCTORY


     
Incense-cedar in Yosemite National Park. Photo by Charles Webber California Academy of Sciences. Photo by Lin1, 2007.
AUTHORSHIP AND CITATION:
Tollefson, Jennifer E. 2008. Calocedrus decurrens. 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:
CALDEC

NRCS PLANT CODE [208]:
CADE27

COMMON NAME:
incense-cedar

TAXONOMY:
The scientific name of incense-cedar is Calocedrus decurrens (Torr.) Florin (Cupressaceae) [51,58,64,96,97,102,103].

SYNONYMS:
Libocedrus decurrens Torr. [161]

LIFE FORM:
Tree

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Information on state-level protected status of plants in the United States is available at Plants Database.

DISTRIBUTION AND OCCURRENCE

SPECIES: Calocedrus decurrens
GENERAL DISTRIBUTION:
Incense-cedar is a native tree in montane forests from Oregon south through California to northern Baja California, Mexico and east to western Nevada [58,64,96,97,103,208]. In Oregon, it extends from the southeastern slopes of Mt Hood east into central Oregon and south along the Cascade Range [7]. Its California distribution includes the Siskiyou, Klamath, North Coast, Cascade, Sierra Nevada, South Coast, Transverse, and Peninsular ranges and the Modoc Plateau [205]. The US Geological Survey provides a distributional map of incense-cedar.

HABITAT TYPES AND PLANT COMMUNITIES:
Incense-cedar is an important component of mixed-conifer forests in southern Oregon, California, and northern Baja California [10,16,71,73,96,103,145,191,207]. Incense-cedar is also common in white fir (Abies concolor) forests at the upper margin of the mixed-conifer zone in southwestern Oregon [10,73] and northern California [182] and giant sequoia (Sequoiadendron giganteum) groves in the Sierra Nevada mixed-conifer zone of California [5,14,33]. In Sierra Nevada mixed-conifer forests it may account for half of the stems in a stand [174]. For information on tree, shrub, and herbaceous species associated with incense-cedar in mixed-conifer forests, see these sources: [10,15,16,71,172].

Incense-cedar is a minor component of the other forest types in which it is found [174]. It occurs with Shasta red fir (Abies magnifica var. shastensis) [11,73] and California red fir (A. magnifica) [15,73]. It grows in Douglas-fir-western hemlock (Pseudotsuga menziesii-Tsuga heterophylla) forests [71,73,160] and in grand fir (A. grandis) forests in southern Oregon [73]. It is a minor component of mixed-evergreen forests in southwestern Oregon and California [16,38,71,73,84,96,97,163] and of redwood (Sequoia sempervirens) forests in north coastal California [71,234].

Incense-cedar occurs with bigcone Douglas-fir (P. macrocarpa) in southern California [140] and with Jeffrey pine (Pinus jeffreyi) and ponderosa pine (Pinus ponderosa var. ponderosa) throughout much of its range [15,87,182]. Incense-cedar and Jeffrey pine are common associates on serpentine soils [10,53,73,101]. On the east side of the Oregon Cascade Range, incense-cedar occurs in dry ponderosa pine forests [200]. On the eastern slope of the Sierra Nevada, it grows with ponderosa pine, Jeffrey pine, sugar pine (P. lambertiana), and white fir [25,45,103,120]. Incense-cedar grows with Oregon white oak (Quercus garryana) [7,73,195] and California black oak (Q. kelloggii) in southern Oregon and California [7,134]. It is a minor associate in canyon live oak (Q. chrysolepis) forests [128] and may also extend into the chaparral zone in California [26,31,185].

Incense-cedar is rarely found in pure stands [114,174].

Vegetation types describing plant communities where incense-cedar is a dominant species are listed below.

Oregon: California: California and Oregon:

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Calocedrus decurrens
 
 

Photo by 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 (e.g., [51,96,97]).

Incense-cedar is an evergreen tree that grows 66 to 187 feet (20-57 m) tall and up to 4 feet (1.2 m) in diameter [48,51,64,70,73,96,97]. In presettlement giant sequoia forests, incense-cedar trees reportedly reached 225 feet (69 m) tall and 12 feet (4 m) in diameter [32,70]. At high elevations and on dry, exposed sites, trees are small and scrubby [174]. Young trees have dense, symmetrical, pyramid-shaped crowns with branches that reach to the ground. Old trees have swollen bases, rapidly tapering trunks, and open, irregular crowns. Very old trees often have dead tops. Trees grow slowly and can live over 500 years [7,48,70,73,174].

Incense-cedar bark is thick, fibrous, furrowed, and ridged [48,51,64,96,97]. The bark is usually 2 to 3 inches (5.0-7.6 cm) thick but may be as thick as 6 to 8 inches (15-20 cm) on old trees. The bark exfoliates into fibrous shreds [46]. Leaves are scale-like, 3 to 14 mm long, and form flat sprays [48,51,64,96,97]. Male cones are terminal on twigs and reach a length of 4 to 7 mm. Female cones develop on the ends of the previous year's growth and reach 0.6 to 1.5 inches (1.4-4 cm) at maturity [48,51,64,96,97,174]. They contain 4 or fewer seeds. Seeds are 8 to 12 mm long and have 2 wings of unequal length [51,64,96,97,174].

Incense-cedar has a well developed root system [174] consisting of widespreading lateral roots and several downward-growing roots. Both lateral roots and taproots branch "profusely". Because new roots commonly branch off at a 45 angle from the parent root, the root system occupies a broad lateral area with depth. Some branches from horizontal lateral roots also grow upward to within 1.2 inches (3 cm) of the soil surface [194].

RAUNKIAER [176] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Incense-cedar reproduces by seed [51,64,96,97].

Pollination: Incense-cedar is wind pollinated [174].

Breeding system: Incense-cedar is monoecious [48,174,205].

Seed production: Incense-cedar seed production varies by tree, year, and location [174]. Trees can produce up to 186,000 seeds/acre (review by [117]). In a heavy seed production year, incense-cedar may produce up to 405,000 seeds/acre [69]. There are approximately 15,000 to 16,000 seeds/lb [4,7,56].

Trees produce abundant seeds every 3 to 6 years. In some years trees produce no seeds [174], (Habeck 1992a, cited in [69]). On the Challenge Experimental Forest in Yuba County, California, incense-cedar produced 10 seed crops in 24 years. Of these, 1 crop was considered "medium to heavy", and 9 crops were considered "very light to light" [139].

Seed dispersal: Incense-cedar seeds are wind dispersed [7]. Because the seeds are light and have a large wing averaging 1 inch (2.5 cm) in length, they fall slowly (5.9 feet (1.8 m)/s in still air) and may be carried great distances by wind [4,65]. In a study of seed dissemination in north-central California, 100% of incense-cedar seeds counted fell within 200 feet (60 m) of the parent tree [137].

Seed banking: No information is available on this topic.

Germination: Incense-cedar seeds germinate well on bare soil and in light litter (review by [117]). Seeds can also germinate on a well-developed duff layer [106]. In a greenhouse experiment, incense-cedar germination was 19% on basalt-derived soil and 18% on sandstone-derived soil [153]. Although germination may be as high as 98% under controlled conditions [174], field germination rates usually vary between 20% and 40% (review by [117]). Incense-cedar germination is "improved" with cold stratification at 37 to 41 F (3-5 C) for 8 weeks [193]. The optimum germination temperature for incense-cedar is 68 F (20 C) (Barton 1930, cited in [20]).

Seedling establishment/growth: Incense-cedar seedlings can establish in shade and in heavy litter or brush cover [7,14,86,106,179]. Seedlings can also establish on mineral soil [7]. In a study of conifer regeneration after logging on the Stanislaus-Tuolumne Experimental Forest, California, incense-cedar germinated best in half shade on bare soil but survived best in half shade on "medium" litter [192].

Incense-cedar seedling density after logging is variable. On the Stanislaus-Tuolumne Experimental Forest, incense-cedar seedling density was 1,080 to 2,190 stems/acre 11 to 12 years after clearcutting [192]. Incense-cedar seedlings were uncommon, however, following clearcutting in Yuba County, California. The seedlings present were concentrated near the shaded edge of the clearcut [138]. On the Challenge Experimental Forest, incense-cedar seedlings were abundant 9 years after shelterwood cutting and absent after clearcutting [183].

Number of incense-cedar seedlings by cutting method 9 years after treatment [183]
Treatment Number of seedlings
Single-tree selection 44
Group selection 16
Shelterwood 470
Seedtree 67
Clearcut 0

Seedling growth is slow [12,192]. Low sunlight and heavy deer browsing are some of the factors that inhibit seedling growth [174]. Incense-cedar often reaches only 3 to 6 inches (8-15 cm) in height after 3 to 5 years. On the Stanislaus-Tuolumne Experimental Forest, the average height of incense-cedar seedlings 12 years after logging was 8 inches (20 cm) [192]. On very dry sites or in dense shade, saplings may only reach 3 feet (0.9 m) in 30 years [7]. The rate of shoot elongation in incense-cedar varies in relation to moisture availability. In a greenhouse experiment, incense-cedar growth rate accelerated after watering and slowed with increasing water stress [90]. Given sufficient water, seedling growth is faster in forest openings than in shade [7].

Incense-cedar seedlings have well-developed root systems [73]. In the first growing season, roots may extend to a depth of 12 inches (30 cm) [65]. Root growth of incense-cedar seedlings after 2 years was greatest in deep, loamy sand at low elevation [194]. Lateral root length was calculated as the average length of the 4 longest lateral roots.

Average root lengths of incense-cedar grown on 3 soil types in the South Umpqua River drainage, Oregon [194].
Soil texture Soil depth (cm) Elevation (m) Taproot length (cm) Lateral root length (cm) Average top:root ratio
Loamy sand 180 210 126.5 130.3 0.45
Loam 100 1,010 79.3 43.2 0.36
Clay loam 160 850 90.3 33.0 0.25

Incense-cedar seedlings are susceptible to mortality from a variety of causes. The average survival rate of first-year incense-cedar seedlings on the Stanislaus-Tuolumne Experimental Forest was 10.3%. Cutworms and drought were the greatest causes of seedling mortality [67].

Causes of mortality and percent of first-year incense-cedar seedlings killed over 8 years [67]
Frost Rodents Insect (cutworms) Fungi Heat Drought Misc.
2.4 4.8 52.0 3.1 0.0 19.8 6.2

Vegetative regeneration: Incense-cedar does not reproduce vegetatively [40,110].

SITE CHARACTERISTICS:
Incense-cedar grows on a wide variety of sites ranging from shaded stream courses to exposed slopes [99]. It grows well on hot, dry sites and commonly occupies an upper canopy position on southwest-facing slopes [174]. It is common on xeric sites in the mixed-conifer zone of southern Oregon and California [34,62,68,73,174,182] and is found on exposed serpentine ridges in the Santa Lucia Range, California [81].

Incense-cedar also occurs on cool, moist sites [73,103,110,172,182], although it often is subdominant to other species on such sites [174]. It occurs in riparian woodlands [133] and is classified as a facultative riparian conifer in the eastern Sierra Nevada (Taylor and Davilla 1985, cited in [89]). In the Santa Lucia Range, incense-cedar is concentrated in deep canyons and shady ravines [81,82]. In the McKenzie River Valley, Oregon, incense-cedar is found on alluvial landforms where the water table remains close to the surface year-round [92]. At the southern extent of mixed-conifer forest in the Sierra San Pedro Mrtir in northern Baja California, incense-cedar occurs almost exclusively on mesic sites including riparian habitats [16,146,150].

Elevation: Incense-cedar occurs between 165 and 6,600 feet (50-2,010 m) in the northern portion of its range and between 3,000 and 9,700 (910-2,960 m) feet at its southern limit. In the Sierra Nevada, incense-cedar grows best between 2,000 and 6,900 feet (610-2,100 m) [174].

Elevational ranges of incense-cedar
Location Elevation
California 980-8,200 feet [96,121,161]
Nevada 5,000-7,000 feet [103]
Baja California, Mexico 3,600-7,900 feet [146,150]

Soils: Incense-cedar grows in many soil types originating from a wide variety of parent rocks including rhyolite, pumice, andesite, diorite, sandstone, shale, basalt, peridotite, serpentinite, granite, and limestone [174]. Incense-cedar is an indicator of serpentine soils in portions of the Klamath Mountains and California's Coast Ranges [80,80,115]. Its ability to extract soil phosphorus and calcium and exclude surplus magnesium allows incense-cedar to grow on soils derived from peridotite or serpentinite [174]. Texture of soils supporting incense-cedar varies from coarse sand to fine clay [174,193]. The best incense-cedar stands are generally found on deep, well-drained, sandy loam and clay loam soils [174]. Incense-cedar grows in pH ranges from strongly acid to nearly neutral [174], although it has a slight affinity toward basic soil conditions [9].

Moisture: Incense-cedar is very drought tolerant [7,206]. It closes its stomata to control water loss on dry sites [95]. Summer precipitation is usually less than 1 inch (25 mm)/month. Incense-cedar can grow on sites that receive as little as 15 inches (380 mm) of annual precipitation [174], but annual precipitation (including snow) varies from 20 to 80 inches (510-2,030 mm) across its range [174,193]. Incense-cedar is intolerant of flooding [228].

Temperature: Incense-cedar is tolerant of heat [62] and somewhat resistant to frost injury [67]. Annual temperature extremes across the range of incense-cedar are -30 F to 118 F (-34 C to 48 C) [174].

SUCCESSIONAL STATUS:
Incense-cedar is shade tolerant [13,59,69,73,167]. Seedlings establish readily in shade, and trees persist in the shaded understory for long periods [14,86,169,179]. In a second-growth mixed-conifer forest in Tulare County, California, growth rings indicated that most of the incense-cedars with a small DBH were nearly as old as the largest individuals [28]. In the northern Oregon Cascade Range, however, incense-cedar is less tolerant of shade than western hemlock, western redcedar (Thuja plicata), and grand fir. It requires occasional disturbance to persist in these stands [7].

In many stands, incense-cedar is an important component of both the understory and the overstory [52,75]. It occupies a "subdominant" crown position in several forest types [174]. Incense-cedar attains canopy tree status by releasing in canopy openings [73,120,216]. Incense-cedar is reported as a late-seral canopy dominant in dry mixed-conifer forests of the western Oregon Cascade Range [57,73], in portions of the white fir zone in southern Oregon [73], and in some mixed-conifer forests [72,184]. Incense-cedar is also a pioneer species in many areas, including high ridges in the Umpqua River drainage and meadow communities in central and southern Oregon [73,86,154,210].

Although recruitment of incense-cedar is not fire-dependent [110], fire does influence its succession. The historic regime of frequent, low-severity fire in mixed-conifer forests favored ponderosa pine and other fire-adapted species over fire-susceptible species such as incense-cedar and white fir [14,35,86,100,219]. After decades of fire exclusion, many mixed-conifer forests in Oregon and California now have dense understories dominated by incense-cedar and other shade-tolerant species [15,16,124,169]. In the long absence of fire or other disturbance, subcanopy incense-cedars eventually grow into the overstory of mixed-conifer forests [72].

SEASONAL DEVELOPMENT:
Incense-cedar has a long growing season that ranges from 91 to 146 days [66,174]. The growing period for incense-cedar at the California Forest and Range Experiment Station in the Sierra Nevada was longer and ended later for incense-cedar than for any of 5 other conifer species studied [66]. Like other members of the cypress family, incense-cedar does not form overwintering buds. Its shoot tips stop growing in the fall and resume growth in the spring [66,90,120]. Seasonal radial growth starts in the spring before height growth [174]. At the California Forest and Range Experiment Station, the 8-year average start date of height growth was 24 May, and the 7-year average start date of radial growth was 15 April [66]. Male cones open and shed pollen in late winter and early spring [7,48,120,174]. Female cones develop in 1 year, maturing by late summer or early fall [48,96,120,174,193]. Seed dispersal begins in late August at low elevations and in October at high elevations and continues into the winter months [65]. On the Challenge Experimental Forest, dissemination of incense-cedar seeds began on 15 September [139].

FIRE ECOLOGY

SPECIES: Calocedrus decurrens
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Mature incense-cedar trees have thick bark that helps protect the tree from low-severity surface fires [7,14,110,132,179,190]. Postfire regeneration is by seeds dispersed in the first postfire year or later by parent trees that survive the fire [110] and by seeds from off-site sources [4,65,197].

Fire regimes: Historic ignition sources in mixed-conifer forests included both lightning and Native Americans [170,171,229,230]. The historic fire regime in mixed-conifer forests was characterized by frequent, low- to moderate-severity surface fires [41,109,112,148,202,230]. Large, severe fires were infrequent in presettlement mixed-conifer forests of Oregon and California [111], although crown fires affecting small areas were probably common (review by [230]). There is some evidence of historic high-severity fires in mixed-conifer forests in the southern Cascade Range and Klamath Mountains [22,24,164,204]. Across the mixed-conifer zone, fire severity varied with slope, aspect, and topographic position [22].

Historic fire-return intervals in mixed-conifer forests ranged from approximately 3 to 30 years [43,74,109,112,127,165,188,202,219,229]. In giant sequoia groves in the Sierra Nevada, Swetnam and others [201] reported mean historic fire-return intervals of 5 to 10 years and a maximum fire-return interval of 20 years. Throughout the mixed-conifer zone, mean fire-return intervals varied with site: mesic and/or sheltered sites burned less often than xeric and/or exposed sites [62,231]. Frequent, low-severity fires killed small trees, including incense-cedar, prevented accumulation of surface fuels, and maintained an open, park-like forest structure [27,40,45,147].

The historic fire regime in mixed-conifer forests favored ponderosa pine and other fire-adapted species over fire-susceptible species such as incense-cedar and white fir [35]. As a result of 19th century logging practices [121,196] and fire exclusion since the early 20th century, however, shade-tolerant incense-cedar and white fir have increased in mixed-conifer forests, often forming dense thickets in the understory [2,3,6,16,27,39,40,60,121,222]. In the absence of fire, incense-cedar and white fir have also proliferated in the understory of giant sequoia groves in the Sierra Nevada [209].

Contemporary fires in mixed-conifer forests are less frequent, larger, and more severe than in the past [41,62,171,189,230]. The buildup of needle litter, duff, dead wood, and understory trees provide ladder fuels and result in high-severity, stand-replacement wildfires [6,27,29,40,69,109,110,122,151,209]. Because small fires are often suppressed, very large fires are more likely to occur during severe fire weather, such as Santa Ana Winds and heat waves [144]. The 1996 Ackerson Fire in Yosemite National Park, California, burned 19,000 acres (7,700 ha) of mixed-conifer forest where dense thickets of incense-cedar and white fir had developed in the understory and created conditions conducive to severe fire [122]. In October 2003, wildfires burned approximately 740,000 acres (300,000 ha) across southern California, including approximately 25,000 acres (10,000 ha) of mixed-conifer forest [77].

Many mixed-conifer forests in the Sierra San Pedro Mrtir in northern Baja California still experience an unmanaged fire regime. The fire regime there is characterized by moderate- to high-severity surface fires that create open, park-like stands of mature trees. Fires may be as large as 12,000 acres (5,000 ha) with relatively long (~50-year) fire-return intervals. The long fire-return interval in the Sierra San Pedro Mrtir is attributed to slow fuel buildup resulting from relatively low photosynthesis rates in evergreen sclerophyllous shrubs and trees and high live fuel moisture in sprouting shrubs [144,151,152].

Incense-cedar grows in a variety of other plant communities, all of which are subject to periodic or frequent fire. For more information, please refer to FEIS reviews of dominant species and the Fire Regime Table below, which provides fire regime information on vegetation communities in which incense-cedar may occur.

Fire regime information on vegetation communities in which incense-cedar may occur. For each community, fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Models [119]. These vegetation models were developed by local experts using available literature, local data, and/or expert opinion as documented in the PDF file linked from each Potential Natural Vegetation Group listed below. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.
Pacific Northwest California Great Basin    
Pacific Northwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northwest Woodland
Oregon white oak-ponderosa pine Replacement 16% 125 100 300
Mixed 2% 900 50  
Surface or low 81% 25 5 30
Pine savannah (ultramafic) Replacement 7% 200 100 300
Surface or low 93% 15 10 20
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Oregon white oak Replacement 3% 275    
Mixed 19% 50    
Surface or low 78% 12.5    
Northwest Forested
Douglas-fir (Willamette Valley foothills) Replacement 18% 150 100 400
Mixed 29% 90 40 150
Surface or low 53% 50 20 80
Oregon coastal tanoak Replacement 10% 250    
Mixed 90% 28 15 40
Ponderosa pine (xeric) Replacement 37% 130    
Mixed 48% 100    
Surface or low 16% 300    
Dry ponderosa pine (mesic) Replacement 5% 125    
Mixed 13% 50    
Surface or low 82% 8    
Douglas-fir-western hemlock (dry mesic) Replacement 25% 300 250 500
Mixed 75% 100 50 150
Douglas-fir-western hemlock (wet mesic) Replacement 71% 400    
Mixed 29% >1,000    
Mixed conifer (southwestern Oregon) Replacement 4% 400    
Mixed 29% 50    
Surface or low 67% 22    
California mixed evergreen (northern California) Replacement 6% 150 100 200
Mixed 29% 33 15 50
Surface or low 64% 15 5 30
Mixed conifer (eastside dry) Replacement 14% 115 70 200
Mixed 21% 75 70 175
Surface or low 64% 25 20 25
Mixed conifer (eastside mesic) Replacement 35% 200    
Mixed 47% 150    
Surface or low 18% 400    
Red fir Replacement 20% 400 150 400
Mixed 80% 100 80 130
California
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
California Shrubland
Montane chaparral Replacement 34% 95    
Mixed 66% 50    
California Woodland
California oak woodlands Replacement 8% 120    
Mixed 2% 500    
Surface or low 91% 10    
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
California Forested
California mixed evergreen Replacement 10% 140 65 700
Mixed 58% 25 10 33
Surface or low 32% 45 7  
Coast redwood Replacement 2% ≥1,000    
Surface or low 98% 20    
Mixed conifer (North Slopes) Replacement 5% 250    
Mixed 7% 200    
Surface or low 88% 15 10 40
Mixed conifer (South Slopes) Replacement 4% 200    
Mixed 16% 50    
Surface or low 80% 10    
Jeffrey pine Replacement 9% 250    
Mixed 17% 130    
Surface or low 74% 30    
Mixed evergreen-bigcone Douglas-fir (southern coastal) Replacement 29% 250    
Mixed 71% 100    
Interior white fir (northeastern California) Replacement 47% 145    
Mixed 32% 210    
Surface or low 21% 325    
Red fir-white fir Replacement 13% 200 125 500
Mixed 36% 70    
Surface or low 51% 50 15 50
Red fir-western white pine Replacement 16% 250    
Mixed 65% 60 25 80
Surface or low 19% 200    
Great Basin
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Basin Shrubland
Montane chaparral Replacement 37% 93    
Mixed 63% 54    
Great Basin Woodland
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Great Basin Forested
Interior ponderosa pine Replacement 5% 161   800
Mixed 10% 80 50 80
Surface or low 86% 9 8 10
*Fire Severities
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [88,118].

Fuels: Fire exclusion has resulted in an increase in understory live fuels in many mixed-conifer forests where incense-cedar seedlings and saplings often form a dense under- and midstory [2,3,3,27,39,109,121]. Due to thin bark, flammable crowns [14,35,69,100,132,219], and branches that often reach to the ground [48], young incense-cedar trees are likely to torch and act as ladder fuels [3]. Incense-cedar seedlings have particularly flammable bark and foliage and are usually totally consumed by fire [7].

In a prescribed fire study in mixed-conifer forest in Yosemite National Park, 1 of the 4 fuel types studied was dominated by incense-cedar seedlings and saplings [213,221]. Prior to burning, mean fuel loads in this fuel type were [213]:

Prefire fine and heavy fuels (g/m). Data are means [213].

Layer Incense-cedar fuel type
surface fuels* 18.0
fresh litter 292.0
weathered litter 431.5
duff 2,830.9
total fine fuel 3,572.4
heavy fuel (>2.5 cm diameter) 1,261.3
*excluding litter

Energy released by the fire in the incense-cedar fuel type was 402.1 kcal/m. Energy release was significantly higher in the incense-cedar fuel type than in types where understory trees were less dense, except where Sierra mountain misery (Chamaebatia foliolosa) dominated the understory; in that type, incense-cedar seedlings were also abundant [221]. For further information on this study, see the Research Project Summary of Van Wagtendonk's [213,214,221] study. This and other studies have demonstrated that prescribed fires can cause some reduction in understory incense-cedar fuels [109,113,130,162,182,221].

POSTFIRE REGENERATION STRATEGY [197]:
Tree without adventitious buds and without a sprouting root crown
Crown residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on-site or off-site seed sources)

FIRE EFFECTS

SPECIES: Calocedrus decurrens

Photo by Walter Roth.


IMMEDIATE FIRE EFFECT ON PLANT:
Young incense-cedar trees are usually killed by fire due to thin bark and flammable crowns [14,35,69,100,132,219]. They often have branches that reach to the ground [48] and are therefore likely to torch [3]. Mature incense-cedars have thick bark and are more fire resistant than young trees [40,100,132,199,211,219]. Mature trees may survive or be killed by fire, depending on the severity of the fire [110,219].

Incense-cedars growing on moist, protected sites are likely to survive fire. Incense-cedars surviving the 1991 Warner Creek Fire on the Willamette National Forest, Oregon, for example, were located on a low slope near a riparian area [175]. Following the August 1977 Marble Cone Fire in Monterey County, California, the only remaining incense-cedar stands were located in deep, moist canyons where the fire was not severe [82].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
The probability of incense-cedar mortality increases with increasing char height and decreases with increasing DBH [178]. Even mature incense-cedars, however, are susceptible to fire damage. The outer bark on incense-cedar is dry, stringy, and deeply furrowed. The thick bark ridges protect the inner bark and cambium from heat injury, but the cambium under the crevices between ridges is easily damaged. As a result, many mature trees in locations subject to past fires have long, narrow fire scars. The susceptibility of incense-cedar to cambial injury from fire makes it a valuable species for tree ring-based fire history studies [226]. According to a 1961 guide for marking fire-damaged timber [225], incense-cedars are likely to survive late-season fires if cambium injury is none to moderate (<25% of cambium killed, little damage above "stump height") and crown mortality is less than 45% to 55%.

PLANT RESPONSE TO FIRE:
Incense-cedar reproduces after fire by seed blown into fire-created canopy gaps by wind. Seeds disperse from surviving parent trees and from off-site sources [50,110,197]. Two years after the October 2003 Cedar Fire in Cuyamaca Rancho State Park, California, thousands of incense-cedar seedlings were observed in a stand with surviving, adult incense-cedars [69]. Severe, stand-replacement fire, however, may kill most seed-bearing incense-cedars. The establishment rate of incense-cedar seedlings within fire-killed stands decreases with distance to the nearest seed source [77]. The stand-replacing portion of the Cedar Fire burned nearly 10,000 acres (4,000 ha) of mixed-conifer forest. In a study of postfire regeneration 4 years later, incense-cedar seedlings were absent from all but 1 of 8 plots. The only plot with incense-cedar seedlings had surviving trees nearby [77]. Incense-cedar seedlings that establish after fire are generally suppressed in the early seral stages by shade-intolerant, fast-growing species such as ponderosa pine and sugar pine. Incense-cedar is shade tolerant, however, and can persist in the understory for many years [45].

Incense-cedar increases in the absence of fire [6,14,100,187]. Historically, frequent, low-severity fire thinned sapling and pole-sized incense-cedars in the understory of mixed-conifer forests [77]. Fire exclusion since the early 1900s has allowed continuous recruitment of incense-cedar and white fir, resulting in dense understory thickets of these shade-tolerant, fire-sensitive species in many mixed-conifer forests [16,19,69,77,109,223]. A study of 68 field quadrats in southern California mixed-conifer forest 60 years after the 1929 to 1934 California Vegetation Type Map Survey showed a 74% increase in stem density, due primarily to a 10-fold increase in incense-cedar and white fir <13 inches (33 cm) DBH [149,151]. In Cuyamaca Rancho State Park, density of pole-sized conifers has increased by 250% since the late 1920s, while old-growth trees have decreased by 40%. Incense-cedar has increased by nearly a factor of 4, largely due to the ingrowth of small trees [77]. In a Sierran mixed-conifer forest, white fir and incense-cedar were 2 to 4 times as important in the sapling layer as in the overstory [17].

Importance values (relative basal area + relative density + relative frequency) of incense-cedar and white fir in 3 size classes in Placer County Big Trees Grove, California [17]
  <3 cm DBH
(saplings)
30-40 cm DBH >40 cm DBH
(overstory trees)
incense-cedar 62 35 27
white fir 132 123 36

Although incense-cedar establishes readily in the absence of fire, it only persists in a stand if fire is absent until young trees are large enough to survive low-severity fire [100].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
No additional information is available on this topic.

FIRE MANAGEMENT CONSIDERATIONS:
Prescribed fire can be used to restore and maintain historic composition and structure in mixed-conifer forests [27,40,78,209,215]. The objectives of prescribed fire in this forest type include reducing the understory density of fire-sensitive, shade-tolerant incense-cedar and white fir [233]. In a mixed-conifer forest in Yosemite National Park, for example, prescribed fire objectives include 50% reduction in total fuel load and mortality of 40% of pole size (1.0-5.9 inches (2.5-15 cm) DBH) incense-cedar and white fir [122].

Numerous case studies illustrate the effectiveness of prescribed fire in reducing understory incense-cedar density. For example, incense-cedar showed greatest mortality among conifers following 4 fire treatments in 1983 and 1984 in mixed-conifer forests of the southern Cascade Range and the northern Sierra Nevada, with mortalities near 100% on some plots. Incense-cedar was "all but eliminated" from the understory by all fire treatments. After a late-spring, moderate-severity treatment, incense-cedar density decreased from 19,967 stems/ha to 67 stems/ha [109]. For further information on this study, see the Research Project Summary describing Kauffman and others's [104,105,107,108,109] research. Mortality following a fall 2002 prescribed fire in the same study area was greatest for incense-cedar in the smallest size class [113].

Postfire percent mortality for incense-cedar in 3 DBH classes [113]
2.5-25 cm DBH 25-51 cm DBH 51-76 cm DBH
27.11 2.11 6.67

Near the Plumas National Forest, California, prescribed fire in a mixed-conifer-California black oak forest with an incense-cedar component successfully reduced fuel loads. When a wildfire burned through the site previously burned under prescription, fire severity and fire suppression costs were less compared to adjacent land where fire had been excluded [155]. For further information on this study, see the Research Paper by Moghaddas [155]. A 1990 fall prescribed fire in the Tharp Creek Watershed of Sequoia National Park, California, produced 16.4% and 17.8% average annual incense-cedar mortality on 2 white fir-mixed conifer sites monitored for 5 years after fire. Mortality was concentrated in the subcanopy [162]. For more information, see the Research Paper by Mutch and Parsons [162].

Prescribed fires in Cuyamaca Rancho State Park demonstrated the effectiveness of spring and late fall underburns in controlling young incense-cedars. Incense-cedar showed significant reductions in both sapling (P<0.02) and seedling densities (P<0.01) after burning [130]. For further information on this study, see the Research Project Summary of Martin and Lathrop's [123,130,131] study. Prescribed burns in mixed-conifer forest in Sequoia and Kings Canyon National Parks reduced the density of small (1-3.5 inches (2.5-9 cm)) incense-cedar by 56% [182].

Typically, prescribed fires in the mixed-conifer zone are conducted in late fall. With relatively low temperatures and high humidity, these late-season fires are generally lower severity than midseason fires and therefore may not kill a sufficient percentage of understory trees to meet restoration objectives. In a series of prescribed fires in Yosemite National Park, total understory incense-cedar basal area was significantly affected by fuel type and fuel moisture. On incense-cedar fuel plots, only plots at the 10% fuel moisture content were sufficiently dry to kill incense-cedars 3 to 10 feet (1-3 m) in height [221]. For further information on this study, see the Research Project Summary of Van Wagtendonk's [213,214,221] study.

A combination of thinning and prescribed burning may more effectively move stands with dense incense-cedar understories toward historic conditions than thinning or prescribed burning alone [166]. Application of 2 or 3 burns may also help to incrementally reduce fuel loadings in such stands [40,109]. Understory thinning may be necessary to allow successful application of prescribed fire in dense stands with high fuel loads [19,29,40,59,60,63,156]. On the Blacks Mountain Experimental Forest in northeastern California, prescribed burning was conducted in October 1959 to kill dense incense-cedar seedlings and saplings beneath a mature stand of ponderosa pine [78]. Due to abundant fuels on the site, the fire was more severe than expected and reached the canopy in some areas. Although the main objective of the fire was met (the incense-cedar understory was reduced from 1,031 to 16 live trees), Gordon [78] recommended against future burning under such extreme fuel conditions.

Even with a combination of fire and thinning treatments, restoring historic composition and structure to mixed-conifer forests after nearly 100 years of fire exclusion may be difficult. Although fire and thinning treatments kill incense-cedar seedlings and saplings, posttreatment seed rain and seedling establishment are often high. Fall prescribed fire and thinning treatments in an unmanaged, old-growth mixed-conifer forest at the Teakettle Experiment Forest, California, resulted in greater incense-cedar sapling reduction in burned vs. unburned treatments (P=0.030) and in thinned vs. unthinned treatments (P=0.036). Incense-cedar seedlings were also significantly reduced in both burned (P=0.0052) and thinned (P=0.0021) sites. Posttreatment seed rain and seedling establishment, however, were up to an order of magnitude higher for incense-cedar and white fir than for pines (Pinus spp.). Under these conditions, stand structure and composition returns to pretreatment conditions within a few years [233]. Regular prescribed fires may be necessary to maintain low density of understory incense-cedars [130]. Incense-cedar seedling establishment is reduced by overstory thinning of large, seed-producing incense-cedar trees. For managers attempting to accelerate old-growth development, however, removal of large incense-cedars may not be a desirable option [233].

For more information on prescribed fire techniques and prescribed fire effects in mixed-conifer forests, see these sources: [27,30,217,218,220].

MANAGEMENT CONSIDERATIONS

SPECIES: Calocedrus decurrens
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Mule deer in California and Nevada browse incense-cedar [99,124,125,181]. A variety of insectivorous birds forage on incense-cedar [37,157,158,159]. White-headed woodpeckers, brown creepers, red-breasted nuthatches, and golden-crowned kinglets are among the species that exhibit the greatest use of incense-cedar [157,159]. Brown creepers forage for arthropods on the surface of incense-cedar bark more than that of any other tree species in mixed-conifer forests on the western slope of the Sierra Nevada. They forage significantly more (P<0.05) on incense-cedar in January and February than in the spring and summer months [1].

Palatability/nutritional value: Incense-cedar seeds contain a pungent resin that makes them unpalatable to rodents [4]. Incense-cedar seeds ranked 7th out of 8 conifer species in order of preference by rodents in the Redwood Mountains giant sequoia grove in California [91]. However, incense-cedar seeds are reportedly a preferred food of dusky-footed woodrats in mixed-conifer forests of Lassen County, California [141].

Cover value: A variety of raptors roost and/or nest in large incense-cedars. The majority of known northern and California spotted owl sites are in mixed-conifer forest [44,79,224]. In the San Bernardino Mountains, California, spotted owl nests were found in 3 incense-cedar trees averaging 46 inches (117 cm) DBH, 131 feet (40 m) tall, and 193 years old. Average nest height was 83 feet (25 m) above ground [85]. Great gray owls are also common in mixed-conifer forest [212,232] and are known to nest in large, broken-topped incense-cedars [23]. In a Klamath County, Oregon, mixed-conifer forest, incense-cedar accounted for 3% of 76 bald eagle roost trees [54].

Small incense-cedar trees create a dense understory that provides cover for small birds, particularly during winter [121,157]. Experimental reduction of incense-cedar density resulted in decreases in the numbers of many bird species; approximately 150 incense-cedar trees <20 cm DBH/ha were required to maintain bird abundance and diversity [157]. Although retention of small incense-cedars is generally contrary to current forest management objectives in mixed-conifer forests, Morrison and others [158] recommend maintaining a high diversity of tree species and size classes throughout the mixed-conifer zone of the Sierra Nevada in order to maintain diverse and abundant bird communities.

VALUE FOR REHABILITATION OF DISTURBED SITES:
Incense-cedar is used for erosion control along road cuts and along streams between 2,000 and 6,000 feet (600-1,800 m) elevation in southern California [99]. Incense-cedar seedlings planted in the spring were more successful than fall plantings in an area disturbed by landslides and avalanches near Lake Tahoe, California [18]. Information on propagation of incense-cedar is provided in Kruckeberg [114].

OTHER USES:
Incense-cedar is an important commercial softwood species [42,93]. Although standing trees are highly susceptible to pocket dry rot (Polyporus amarus), products manufactured from incense-cedar wood are extremely durable and decay resistant [42]. Incense-cedar wood is used for many products including lumber, fence posts, railroad ties, venetian blinds, greenhouse benches, siding, decking, cedar chests, and shingles [51,64,103,121]. It is the major source of pencil stock in the United States [64,103]. Incense-cedar is also widely grown as an ornamental tree [64,97,114].

Incense-cedar was used by the Cahuilla of California to construct conical-shaped bark houses that were used for temporary shelter during acorn gathering times in late fall. In some areas, incense-cedar slabs were used in more permanent house construction [21]. Incense-cedar leaves were used by Native Americans of Mendocino County, California, in the process of leaching acorn meal and in a decoction for relieving stomach upset. Small limbs were sometimes used for bows [47].

OTHER MANAGEMENT CONSIDERATIONS:
Incense-cedar is highly susceptible to pocket dry rot, particularly in the mild, western portions of incense-cedar's range [42,227]. Pocket dry rot is most prevalent in trees >150 years old and commonly enters the tree through fire scars and the wounds caused by broken branches [227]. An estimated 81% of pocket dry rot infections enter through fire scars [36]. In parts of the Sierra Nevada, 75% to 100% of mature trees are infected (Bega 1978, cited in [174]). In a 1958 Forest Service, US Department of Agriculture publication, Wagener and Bega [227] estimated that pocket dry rot resulted in more than 36% cull of the standing volume of incense-cedar in California. Cull may be as high as 77% for "overmature" trees [174]. During logging operations in the late 1800s in Sierra Nevada mixed-conifer forests, many incense-cedars were left standing as seed trees due to pocket dry rot that made their lumber virtually worthless. These trees self-seeded, resulting in a dense understory of incense-cedar after pines resumed canopy dominance in cutover stands. Wildfire exclusion after the turn of the century further favored the incense-cedars, allowing them to persist in dense understories [121,196].

Incense-cedar is susceptible to a variety of other pathogens including annosus root disease (Heterobasidion annosum) [55,121,129,180], the trunk rot fungus Oligoporus amarus [15], incense-cedar rust (Gymnosporangium libocedri) [15,168], and the western conifer seed bug (Leptoglossus occidentalis) [61]. Incense-cedar mistletoe (Phoradendron libocedri) is common in incense-cedar crowns [76]. Incense-cedar is occasionally infested with mountain pine beetles (Dendroctonus ponderosae), but the beetles rarely produce broods in incense-cedar [94].

Incense-cedar is less susceptible to ozone-induced injury than other western conifers [143]. In the San Bernardino Mountains, areas of mixed-conifer forest may eventually shift in dominance to incense-cedar as ozone-susceptible ponderosa pine declines (McBride 1985, cited in [83]).

Information on the effects of herbicides on incense-cedar is provided in Conard and Emmingham [49].

Calocedrus decurrens: REFERENCES


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