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Incense-cedar in Yosemite National Park. Photo by Charles Webber © California Academy of Sciences. | Photo by Liné1, 2007. |
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:![]() |
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Photo by Charles Webber © California Academy of Sciences. |
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: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 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 Mártir 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: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: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 Mártir 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 Mártir 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. | ||||||||||
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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) |
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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) |
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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) |
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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/m²). Data are means [213]. |
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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 |
Energy released by the fire in the incense-cedar fuel type was 402.1 kcal/m². 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]:![]() |
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Photo by Walter Roth. |
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: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: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].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 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 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]).
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