Artemisia californica



INTRODUCTORY


  James L. Reveal @ USDA-NRCS PLANTS Database
AUTHORSHIP AND CITATION:
Hauser, A. Scott. 2006. Artemisia californica. 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:
ARTCAL

SYNONYMS:
Artemisia californica var. californica Munz [155,156]

NRCS PLANT CODE [191]:
ARCA11

COMMON NAMES:
California sagebrush
coastal sagebrush
California sagewort
California mugwort

TAXONOMY:
The currently accepted scientific name of California sagebrush is Artemisia californica Less. (Asteraceae) [60,94,107,210].

LIFE FORM:
Shrub

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: Artemisia californica
GENERAL DISTRIBUTION:
California sagebrush is endemic to California [60,107,155,156] and Baja California [118,204,210]. The distribution of California sagebrush is categorized within 4 major floristic associations. The floristic associations and their ranges are as follows: 1) Diablan: found from San Francisco region south to northern Santa Barbara County; 2) Venturan: a coastal group found from northern Santa Barbara County south through coastal Los Angeles County; 3) Riversidian: a cismontane inland group found from inland Los Angeles County, western Riverside County, and inland San Diego County; and 4) Diegan: a Baja-influenced group found from Orange County and coastal San Diego County south into northwestern Baja California [13,204]. California sagebrush distribution and occurrence is greatest in the Riversidian floristic association [202]. California sagebrush also occurs on some of the Channel Islands (Santa Clemente, Santa Catalina, Santa Cruz, San Miguel, San Nicholas, and Santa Rosa ) [146,155,169,205]. In Baja California, California sagebrush extends 200 miles (325 km) south of the Mexican border to approximately El Rosario [5,51,118,204]. Plants Database provides a distributional map of California sagebrush.

ECOSYSTEMS [64]:
FRES28 Western hardwoods
FRES34 Chaparral-mountain shrub
FRES35 Pinyon-juniper
FRES42 Annual grasslands

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES
CA

MEXICO
B.C.N.

BLM PHYSIOGRAPHIC REGIONS [20]:
3 Southern Pacific Border

KUCHLER [119] PLANT ASSOCIATIONS:
K023 Juniper-pinyon woodland
K030 California oakwoods
K033 Chaparral
K035 Coastal sagebrush
K036 Mosaic of K030 and K035
K048 California steppe

SAF COVER TYPES [59]:
239 Pinyon-juniper
250 Blue oak-foothills pine
255 California coast live oak

SRM (RANGELAND) COVER TYPES [179]:
201 Blue oak woodland
202 Coast live oak woodland
204 North coastal shrub
205 Coastal sage shrub
206 Chamise chaparral
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
412 Juniper-pinyon woodland

HABITAT TYPES AND PLANT COMMUNITIES:
California sagebrush is the most important and widely distributed plant in the coastal sage scrub or "soft chaparral" community type [41,86,103,117,188,202]. It also occurs with limited distribution in "hard chaparral" communities dominated or codominated by chamise (Adenostoma fasciculatum), bigpod ceanothus (Ceanothus megacarpus), hoaryleaf ceanothus (C. crassifolius), California scrub oak (Quercus dumosa), curlleaf mountain-mahogany (Cercocarpus ledifolius), sugar sumac (Rhus ovata), toyon (Heteromeles arbutifolia), and hollyleaf cherry (Prunus ilicifolia) [62,70,86,140,188]. In the Great Basin sage scrub community of the San Bernardino Mountains, California sagebrush is codominant with rubber rabbitbrush(Chrysothamnus nauseosus) and antelope bitterbrush (Purshia tridentata) [144]. In plant communities adjacent to coastal sage scrub, overstory associates of California sagebrush include bigcone Douglas-fir (Pseudotsuga macrocarpa), Monterey pine (Pinus radiata), bishop pine (P. muricata), singleleaf pinyon (P. monophylla), California juniper (Juniperus californica), and Tecate cypress (Cupressus forbesii) [10,26,99,103,134,135,140,150,169,188,194,196].

In 1990 and 1991, Davis and others [51] used satellite imagery, air photos, existing vegetation maps, and field data to measure vegetation cover for 8,260,000 acres (3,383,160 ha) within the California Floristic Province. The researchers classified California sagebrush as codominant over 629,600 acres (254,800 ha), of which 356,749 acres (144,371 ha) is typed as coastal sage scrub [51].

California sagebrush is listed as a dominant or codominant species in the following locations and vegetation classifications:


BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Artemisia californica
 

 

James L. Reveal @ USDA-NRCS PLANTS Database
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., [60,94,155,156,210]).

California sagebrush is a drought-deciduous subshrub [14,21,30,52,63,93,168,210] with several to numerous stems arising from the root crown [79]. It grows to a height of 2 to 5 feet (0.5-1.5 m) and is the tallest shrub in the coastal sage scrub community [146,155,156,210]. Lower branches are woody and generally do not exceed 0.2 inch (50 mm) in diameter [129]. California sagebrush branches support numerous leaves from 0.8 to 2 inches (2-5 cm) long and 0.5 to 1 mm wide [79,155,156]. Leaves are seasonally dimorphic [203]. Leaves attached to the main branches or stems are slightly larger and appear early in the growing season. Most of the larger leaves remain on the stem during the dry season, although they may wilt [68,70,204]. On lower branches, side-shoots develop from the leaf axils of larger leaves. The side-shoots develop smaller, persistent leaves [70,72]. The smaller leaves remain wilted for long periods of time under water stress and rehydrate within hours of rainfall [204]. Due to the presence of terpenes, leaves are highly aromatic [93,209]. The inflorescence, 4 to 12 inches (10-30 cm) long and 0.4 to 4 inches (1-10 cm) broad [79], produces 20 to 50 flower heads, with each head containing 15 to 30 disk florets [79,146]. Inflorescences also produce 6 to 10 pistillate flowers [60,94]. California sagebrush flowers are normally yellowish or brownish in color, but along desert borders they are commonly red [79]. The fruit, an achene, is extremely small and lightweight (60 g/seed) [15,54,79,146].

Allelopathy: The essential oil from California sagebrush contains 5 toxic terpenes. It has been suggested that the release of terpenes by California sagebrush contributes to the relative lack of vegetation under and adjacent to the shrub [82,121,213]. During the 1st rains of December, the leaf drip from California sagebrush is toxic. The rain leaches toxins from the leaves and litter that is absorbed by the soil, adding to toxins previously deposited by volatilization during the dry season [116].

Community structure and productivity: In a coastal sage scrub community in the Santa Monica Mountains, California sagebrush and black sage comprise approximately 75% of the community cover. The mean annual aboveground live biomass, mean annual aboveground primary production, and litterfall of the coastal sage scrub community are 925 g/m, 355 g/m, and 199 g/m, respectively [68]. Similar research in the Santa Monica Mountains assessed the biomass and production of California sagebrush in stands burned 22 years prior to the measurements. California sagebrush peak aboveground biomass (June) and net annual production are presented in the following table [72]:

  Foliage Wood Dead Twigs Inflorescences Twigs Total
Peak aboveground biomass (g/m) 51.4 377.4 101.4 ----* ---- 530.2
Net annual production (g/m/yr) 31.3 ---- ---- 3.1 20.6 55.0
*No data

Photosynthesis/transpiration: California sagebrush plants have a thin leaf cuticle and numerous stomata that allow for a high photosynthetic rate in response to water availability. Consequently, California sagebrush has a higher transpiration rate and a longer period of water stress than sclerophyllous species with thick leaves and deep roots [57,139,146].

Roots: California sagebrush plants have fragile, fibrous roots that penetrate shallowly into the soil [9,30,92,139,146,163]. The shallow root system allows for rapid soil moisture absorption and growth at the beginning of the rainy season [56,139,146].

Adventitious rooting was observed within California sagebrush communities at Starr Ranch, Orange County. The development of adventitious roots occurred at the basal portions of their stems, an area covered with soil from erosion or silt deposits. In areas of packed soil, adventitious roots did not occur in California sagebrush. Of 98 California sagebrush plants sampled, approximately 10% exhibited adventitious roots [122].

RAUNKIAER [172] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
California sagebrush regenerates by seed [54,56,57,111,141,146,150,152] and sprouting from the root crown [45,70,78,108,125,126,127,128,146,204] following disturbances such as fire (see Plant Response to Fire).

Pollination: California sagebrush is wind pollinated [146].

Breeding system: California sagebrush has perfect flowers [60,210].

Seed production: California sagebrush plants produce an "abundant" number of seeds annually. In southeastern Orange County, the average seed production of individual California sagebrush shrubs is usually greatest on south-facing slopes. DeSimone and Zedler [54] sampled the seed production of 16 California sagebrush plants on north- and south-facing slopes from February 1995 to May 1996, and sampled on south-facing slopes from fall 1996 to winter 1997. The average seed production per shrub (no. of seedssx) on north-facing slopes from February 1995 to May 1996 was 21,35711,163, compared to 43,63819,671 on south-facing slopes. Seed production on south-facing sites from fall 1996 to winter 1997 was 38,2289,012.

Seed dispersal: California sagebrush seeds are primarily wind dispersed [54,56,57,141,146,150]. Animal dispersion is noted in the literature as a secondary method of dispersal [56,57]. The seeds of California sagebrush are primarily dispersed during autumn [108]. In southeastern Orange County, the dispersal of California sagebrush seeds in an annual grassland, an ecotone (10%-20% shrub cover), and a coastal sage scrub community was measured from February 1995 to May 1996. Seed counts were taken on both north- and south-facing slopes. In the grassland, California sagebrush seeds were dispersed at maximum 15 feet (5 m) from the isolated shrubs. In total, most seeds were found on south-facing slopes, though seed numbers were similar in coastal sage scrub. The following table provides California sagebrush seed counts (no. of seeds/msx) for north- and south-facing slopes in the 3 communities [54]:

Aspect Grassland* (n = 10) Ecotone (n = 4) Coastal sage scrub (n = 4)
North-facing 6961 28474 73475
South-facing 849435 1,103357 730224
*Contained isolated California sagebrush plants

Seed banking: California sagebrush utilizes a seed bank [198]. As of 2006, the literature did not describe the longevity of California sagebrush seeds in the seed bank. Due to the "steady" germination of soil-stored seeds, California sagebrush often fails to develop a seed bank adequate for the recruitment of large numbers of seedlings during the 1st postdisturbance growing season (review by [108]).

In southeastern Orange County, germinable soil seed bank densities of California sagebrush were measured in spring 1995 and 1996 on north- and south-facing slopes of grassland, ecotone, and coastal sage scrub communities. The following table provides California sagebrush germinable soil seed bank densities (no. of seeds/msx) for north- and south-facing slopes in the 3 communities [54]:

Aspect Grassland (n = 10) Ecotone (n = 4) Coastal sage scrub (n = 10)
North-facing 5831 5019 22272
South-facing 4230 20098 584208

Germination: California sagebrush seed germination is induced by several factors. Light, charate, or heat and charate can break seed dormancy [17,54,56,57,108,109,110,146,161]. Seeds near the soil surface may germinate readily with the addition of adequate moisture that occurs with the onset of fall and winter rains [17,54,56,57,110,146]. Seed germination is also favored by cool soil temperatures [56,57] and a day/night temperature regime of approximately 70 F/40 F (20 C/6 C) [17,198]. Keeley [108] showed the rate of germination by seeds is also influenced by heat and charate (see below).

Fire-induced germination: Seeds can be stimulated to germinate by heat and/or chamise charate [17,108,109,110,161]. Two studies on the effects of heat and charate on California sagebrush seeds buried in the soil at and below light penetration level are presented below.

Keeley [108] investigated the effect light, darkness, temperature, and charate have on California sagebrush seed germination. Germination trials were conducted on filter paper and in soil. California sagebrush seeds germinated significantly (p<0.001) better in the soil treatment, and the following results are from the seeds germinated in soil. Temperature treatments were a control, 158 F (70 C) for 1 hour; 212 F (100 C) for 5 minutes; and 248 F (120 C) for 5 minutes. One set of seeds received a 0.50g application of chamise charate prior to temperature treatments. Following charate and temperature treatments, the seeds were stratified for 1 month at 41 F (5 C) and then incubated in the light or dark treatment for 3 weeks at 73 F (23 C). Both light and charate had a significant (p<0.05) positive effect on the germination of California sagebrush seeds. Seeds with charate application and subjected to light germinated significantly (p<0.05) better than treatment seeds subjected to only darkness. Seeds subjected to darkness and charate treatment germinated significantly (p<0.05) better than seeds subjected to only darkness. Within the light treatment, the only significant (p<0.05) difference was the germination rate of charated seeds subjected to the 212 F (100 C) for 5 minutes treatment (81%) and noncharated seeds subjected to the same temperature treatment. When assessing the effect temperature alone has on germination, there was no significance found. Greater than 75% of the California sagebrush seeds germination occurred by the 1st week of the 73 F (23 C) incubation period (reviews by [108,109,110]).

Treatment Percentage germination
Light Dark
Control 158 F
(1 hour)
212 F
(5 min)
248 F
(5 min)
Control 158 F
(1 hour)
212 F
(5 min)
248 F
(5 min)
Control 73 56 47 56 0 10 3 0
Charate 78 80 81 87 62 49 64 50

Seed banked California sagebrush seeds that were subjected to artificial fire conditions produced substantially more germinants than seeds exposed to prescribed fire in the field. California sagebrush seeds obtained from a prefire soil seed bank in a chamise-dominated chaparral community in central California during fall 1989 were treated with heat (212 F (100 C) for 5 minutes) and charate (2 tablespoons of charred chamise/pot) and germinated in a greenhouse. The heat and charate treatment produced a mean of 259.6 California sagebrush germinants/m. Shortly after seeds were collected for the preceding experiment, the site was burned for the 1st time in approximately 50 years. The day after the fire, the researcher collected seeds from the seed bank and planted them in a greenhouse with no further treatment. The rate of germination in the greenhouse (4.4 germinants/m) far exceeded natural seed germination in the field (0.04 germinant/m) following fire but was less than germination from seeds treated with heat and charate [161].

Seedling establishment/growth: California sagebrush seedlings emerge during the rainy season (November-April), and most growth is completed by May with the onset of the dry season [146]. Seedling cover increases for a time after the growth and flowering of young plants. As California sagebrush stands mature and the canopy closes, seedling establishment becomes "poor" [154]. California sagebrush seedlings suffer high rates of mortality due to small animal herbivory [146] and may suffer mortality or inhibited growth due to California sagebrush allelopathy [154]. In areas invaded by annual grasses, California sagebrush seedlings are "outcompeted" for soil moisture, and the rate of survival in their 1st year is "very" low [54]. California sagebrush seedling emergence after fire is "variable" and generally "low". California sagebrush seedlings commonly appear during postfire year 2 from seeds of sprouting plants or seeds dispersed on-site [146,204].

California sagebrush seedling recruitment appears to occur in gaps without fire. In southeastern Orange County, California sagebrush seedling density in 3 communities (annual grassland, ecotone, and coastal sage scrub), on 3 aspects (south-, north-, and west-facing), and in vegetation gaps and under vegetation were measured to assess seedling recruitment between fires. Seedling measurements occurred during March 1995 and 1996 on sites fire-free for 17 years. In the 3 communities and 3 aspects, California sagebrush seedling density was greatest in vegetation gaps. California sagebrush seedlings were significantly (p<0.001) associated with gaps in mature vegetation in coastal sagebrush communities. Vegetation gaps in grasslands were pocket gopher mounds, gaps in ecotones were bare areas between perennial grasses, and gaps in scrub were areas devoid of shrub canopy. California sagebrush seedling density (no. of seedlings/m) on the various sites is presented below [54]:

Community South-facing North-facing West-facing
Gaps Under vegetation Gaps Under vegetation Gaps Under vegetation
Grassland 0.20 0.01 0 0 0 0
Ecotone 5.47 3.12 0 0 0.59 0
Coastal sage scrub 6.60 2.98 0.74 0.11 1.05 0.68

As further evidence that California sagebrush recruitment takes place between fires, juvenile (<20 inches (50 cm) in height and nonreproductive) shrubs were found in all 3 communities and aspects. Recruitment was greatest on west-facing slopes. The density (no. of juveniles/m) of juvenile California sagebrush shrubs is presented below [54]:

Community South-facing North-facing West-facing
Grassland 0.05 0.04 0.42
Ecotone 0.24 0.66 1.47
Scrub 0.36 0.04 1.23

In the coastal sage scrub and grassland communities, California sagebrush seedling growth was measured at the end of May in 1995 and 1996 on south-facing slopes in coastal sage scrub and annual grassland sites. The rate of growth of California sagebrush seedlings in the grassland community was much greater than growth in the coastal sage scrub community. The height (cms) of California sagebrush seedlings in the 2 sites is presented below [54]:

Site 1995 1996
Coastal sage scrub 3.712.69 5.003.06
Grassland 18.672.08 44.007.94

California sagebrush density can depend on winter precipitation. On 16 September 1935 a fire burned in Frankish Canyon at the western edge of San Bernardino County. The number of California sagebrush seedlings/milacre surviving after fire and their average height were measured 8 times over a 17-year period following the fire. The researchers attribute the high number of California sagebrush plants/milacre in postfire year 17 to above normal precipitation during the winter of 1951 to 1952 [101].

  Years after fire
1 2 3 4 5 7 10 17
Surviving
seedlings/milacre
3.2 2.2 2.0 2.2 2.0 4.9 4.1 14.8
Average height (inches) 2.7 9.1 16.3 19.3 21.1 24.1 29.9 ----

Vegetative regeneration: Following damage to aboveground portions of the plant, California sagebrush sprouts from the root crown. Vegetative sprouting is the most common regeneration method following fire [45,70,78,108,125,126,127,146,204]. To study vegetative regeneration of California sagebrush in the absence of fire, Malanson and Westman [130] cut 50 mature California sagebrush shrubs to ground level in the Santa Monica Mountains on a site unburned for 11 years. Two years after cutting, 80% of the shrubs had sprouted [130].

SITE CHARACTERISTICS:
California sagebrush is primarily associated with coastal sage scrub sites, but has limited occurrence in chaparral communities. It is best developed on the coastal side of mountains below chaparral [85,88].

Air pollution: California sagebrush is sensitive to sulphur dioxide pollution. A significant (p<0.01) decrease in California sagebrush foliar cover occurred in shrubs located from 1,600 to 4,900 feet (500-1,500 m) downwind from an oil refinery's sulphur dioxide stacks near Santa Maria, California. Annual sulphur dioxide concentrations (25-year period) emitted from the oil refinery range from 0.09 to 0.17 ppm [171]. Further studies have been conducted on the response of California sagebrush to air pollution in the Santa Monica Mountains near Los Angeles. Where above-average levels of sulphur dioxide and ozone persist, visible foliar damage (fasciated stems) was present [206].

Aspect: California sagebrush plants located on north-facing sites of coastal sage scrub in southeastern Orange County are significantly (p<0.001) taller and have significantly (p<0.002) greater aerial cover/shrub/m than plants on south-facing sites [54].

Aspect Height (msx) Aerial cover/shrub/msx
North 1.340.04 1.040.10
South 1.110.05 0.590.09

At 120 sites in southern California, the frequency (%) of California sagebrush on 2 aspects and 3 slopes was measured. The averages are presented in the table below [118]:

Community type Aspect Slope
North-facing South-facing 0-5 6-15 16+
California sagebrush 9 12 8 17 8
California sagebrush-eastern Mojave buckwheat-white sage 13 16 23 17 12
California sagebrush-coyote bush-basin wildrye 9 3 8 6 6
San Luis purple sage-California sagebrush 30+ 10 8 17 22
California brittlebush-California sagebrush 0 10 0 3 8

Climate: California sagebrush primarily occurs in the mediterranean climate zone, which is characterized by hot, dry summers and mild, moist winters [62,118]. California sagebrush has a bimodal distribution in that it can occur in both the coastal plain, maritime climate and the hot interior of the coastal valleys [40,84,93]. Average yearly precipitation ranges from 10 to 18 inches (250-450 mm) [18,30,40,50,54,118,149,166,207], and approximately 90% of the precipitation falls between November and April [18,40,50,118].

In a modeling experiment, Malanson and others [131] estimate that optimal California sagebrush growth occurs when precipitation ranges from 4 to 10 inches (100-250 mm), the coldest winter temperatures range from 50 F to 54 F (9-12 C), and the warmest summer temperatures range from 75 F to 81 F (24-27 C). Extensive climatic data from the San Gabriel Mountains where California sagebrush is dominant can be found in the review by Miller [138].

Community zonation: Where California sagebrush encroaches into grasslands, the area surrounding California sagebrush generally looks bare. The bare zone, often 3 to 6 feet (1-2 m) wide, has been attributed to allelopathy, animal influences, and soil moisture [16,57,98,116,116,118,150,153,154,173,209].

Though appearing bare in comparison to the grassland and shrub communities, the bare zone is actually comprised of short herbs approximately 1 inch (3 cm) tall. Bare-looking zones generally occur around large, dense shrub assemblages, not where individual or scattered shrubs occur [80,81,116].

At 2 sites, 1 near La Purisima Mission, the other at the base of the Santa Ynez Mountains, Halligan [80] researched possible causes of bare zones surrounding California sagebrush stands. To test the hypothesis that herbivory and granivory causes bare zones between grasslands and shrub zones, the researcher fenced in a 3,400 ft (320 m) area encompassing grassland, the bare zone, and shrubland. Following 1 year of protection, the grassland had invaded 20 inches (40 cm) into the border zone. During the same period in an unprotected area, the grassland regressed from the shrub stand, leaving the border zone 30 inches (70 cm) wider. Within the new border zone, an abundance of brush rabbit feces was present and short animal trails were noticeable, suggesting that small mammals play a part in the creation of bare zones. The research further concluded that California sagebrush allelopathy was a minor contributor to the creation of bare zones. The growth of 2 common herbs, coast tarweed (Madia sativa) and smooth catsear (Hypochaeris glabra), planted under California sagebrush and exposed to the shrub's volatiles, was significantly inhibited (p<0.05). However, 2 common grass species (ripgut brome (Bromus diandrus ssp. rigidus) and rattail fescue (Vulpia myuros)) were not significantly affected by California sagebrush allelopathy. Halligan also found that soil moisture, rainfall, and throughfall of precipitation in the shrub zone are more variable than in grasslands, which also plays a minor part in the presence of bare zones. Further research is available on bare areas associated with California sagebrush [81,83].

Elevation: California sagebrush occurs from 0 to 3,000 feet (0-800 m) but is most common from 500 to 1,500 feet (200-460 m) [85,88,94,155,156]. Its presence is scattered and isolated above 1,500 feet (460 m), where it generally occurs in chaparral belts on shallow soils [85,88,114].

The percent frequency of California sagebrush-dominated communities at different elevations was measured at 120 sites in southern California. The findings are presented in the table below [118]:

Elevation (% frequency)

Community type 0-500 feet 500-1,000 feet 1,000-1,500 feet 1,500-2,000 feet 2000+ feet
Monotypic 0 0 13 19 23
California sagebrush-eastern Mojave buckwheat-white sage 0 18 16 23 23
California sagebrush-coyote bush-basin wildrye 7 12 6 0 8
San Luis purple sage-California sagebrush 25 47 16 0 23
California brittlebush-California sagebrush 21 0 0 3 0

Floodplains: California sagebrush may appear in riparian and floodzones [90,115,181]. Due to the lack of perennial water, several floodplain systems in southern California are composed of unique scrub vegetation rather than the more common riparian woodlands. Of the 10 major floodplain scrub communities, California sagebrush occurs in 4 (San Sevaine, Cucamonga, San Antonio, and San Gabriel). Floodplain scrub vegetation is periodically subjected to severe flooding, erosion, nutrient-poor substrates, and the presence of subsurface moisture. Floodplain scrub communities are found exclusively on the coastal side of major mountain ranges in southern California [90].

Litter: California sagebrush tends to occupy and dominate in sites with a "low" litter mass [202].

Soil: California sagebrush occurs on virtually all soil types, excluding serpentine [42,146]. It has no strong soil type preference, making it the most common species in coastal sage scrub communities [57]. It is the only dominant plant in coastal sage scrub communities that does not exhibit a significant (p<0.01) substrate preference [202]. Soil/substrate types associated with California sagebrush include loam to clayey loams [15,78], sandy loams, and loamy sands. The soils often containing large amounts of gravel, are typically thin and undeveloped, and have little ability to retain nutrients and water [1]. Extensive soil data from a California sagebrush community on Banner Ridge, San Diego County, are presented in a review by Bradbury [29] and from boundary areas between annual grasslands and coastal sage scrub in the review by Hobbs [98].

Transpiration and photosynthesis: In coastal sage scrub communities of the Santa Monica Mountains, California sagebrush has the lowest maximum rate of transpiration and photosynthesis of the dominant plants. The maximum transpiration (g H2O/dm/hr) and photosynthesis (mg CO2/dm/hr) rates for California sagebrush in 1968 were 0.67 and 4.02, respectively [93].

SUCCESSIONAL STATUS:
California sagebrush is adapted to disturbed sites [61,215], is shade intolerant [54,54,61], and occurs across the successional gradient [90]. It is a pioneer species [34,61] in primary and secondary succession and persists as a dominant species through late seral stages in coastal sage scrub communities [18,36,44,90,138,215]. Hall and Clements [79] describe California sagebrush as typically a late successional dominant, but often seral to the chaparral association. This likely occurs in the higher elevation chaparral belt where California sagebrush is a postdisturbance colonizer on steep and/or rocky xeric sites [175]. Disturbances common at these sites include long-term overgrazing or changes in historic fire regimes [195].

Researchers have suggested that various successional progressions where California sagebrush and other dominant coastal sage scrub species occur are influenced by dynamic community boundaries, intervals between fires and other disturbances, and competition between species for environmental resources following disturbance [5,71]. The following section presents research on the effect different disturbance mechanisms have on coastal sage scrub succession.

Anthropogenic disturbance: Within 2 years following the construction of an oil pipeline in the Purisima Hills, Santa Barbara County, California sagebrush had naturally reestablished on the right-of-way. Two years after initiation of growth, shrubs were from 12 to 22 inches (30-55 cm) tall and 14 to 20 inches (35-50 cm) in diameter [36].

In 1985, California sagebrush, along with white sage, black sage, eastern Mojave buckwheat, and brittle bush were outplanted on unamended soil (Arlington loam) in a native plant garden at the Forest Fire Laboratory in Riverside. Inadvertently, the aboveground biomass of the plants was mechanically removed in 1991. Four years later, California sagebrush had regenerated. The density, relative cover, and biomass of California sagebrush in October 1995 were 8.04/m, 68.5%, and 238 g/m, respectively. Given that no coastal sage scrub parcels were in close proximity to the mechanically cleared plots, reestablishment was from on-site seed and root crown sprouting [158].

In San Diego County, severe disturbance (construction, landfill operations, heavy-vehicle activity, tillage, and soil excavation) caused the near extirpation of California sagebrush and other coastal sage scrub species, replacing them with the herbaceous species rattail fescue, brome (Bromus spp.), oatgrass (Danthonia spp.), longbeak stork's bill (Erodium botrys), and smooth catsear (Hypochaeris glabra). While California sagebrush, white sage, chamise, and eastern Mojave buckwheat were the most abundant shrub species on undisturbed sites, these shrub species combined never made up more than 5% of total cover, even when nearby seed sources were available, on sites disturbed 1 to 43 years prior to the study [185].

Community transition: Frequent disturbance in coastal sage scrub communities has strong influence on community structure and transition. Keeley and Keeley [114] hypothesize that recurrent disturbance in coastal sage scrub communities facilitates successional replacement by annual grasslands. Horton [100] suggests that in the absence of disturbance, coastal sage scrub is replaced by coast live oak. Utilizing aerial photography, Callaway and Davis [37] measured transition rates among annual grassland, coastal sage scrub (codominated by California sagebrush), chaparral, and coast live oak woodland communities in Gaviota State Park from 1947 to 1989. In unburned and ungrazed plots, a transition from grassland to coastal sage scrub occurred at a rate of 0.69%/year, from coastal sage scrub to coast live oak woodland at a rate of 0.30%/year, and from coastal sage scrub to chaparral at a rate of 0.11%/year.

Fire: California sagebrush is adapted to burned sites and is an early successional species following fire [18,47,61,85,143,182]. Recovery of coastal sage scrub species from fires that occur at natural, decades-long intervals is rapid. During postfire years 1 and 2, a flush of native annual forbs (sometimes mixed with exotic grasses and forbs) is common, followed by recovery of native shrubs via the seed bank, off-site seed sources, sprouting, and seeds produced by the sprouting shrubs [6,127].

Whereas most California sagebrush regeneration following fire is primarily via sprouting [108,160,204] and relatively rapid, Myers and Ellestrand [157] found that postfire growth at an inland sage scrub site (Motte Rimrock Reserve, Riverside County) was via seedlings and delayed. At the end of postfire year 2, sage scrub (California sagebrush, eastern Mojave buckwheat, brittle bush, and black sage) cover was only 4% of cover on adjacent unburned sites. By the end of postfire year 4, shrub cover was roughly 50% of cover on unburned sites.

California sagebrush seedlings are also pervasive in recently burned chaparral sites but become absent as the stands mature [50,87]. Several studies in the Santa Monica and San Gabriel mountains have found that California sagebrush establishes in chaparral following fire, but over time it is replaced by species that were dominant before fire [71,86,101,103]. At the boundary between chamise chaparral and coastal sage scrub communities (around 3,000 feet (900 m) elevation), a dynamic relationship occurs when fire frequencies are increased. Following fire disturbances in chamise chaparral, coastal sage scrub species often invade the site. The resulting vegetation mosaic is often called the coastal sage-chaparral "subclimax". Species diversity in the coastal sage-chaparral "subclimax" community is often low and common species include eastern Mojave buckwheat, California sagebrush, white sage, black sage, golden-yarrow (Eriophyllum confertiflorum), cudweed (Gnaphalium spp.), and sawtooth bristleweed (Hazardia squarrosa var. grindelioides) [87,88]. (For further information, see Plant Response to Fire.)

Nitrogen deposition: California sagebrush populations have been declining over the past 60 years or more in coastal sage scrub vegetation of southern California. California sagebrush is steadily being displaced by Mediterranean annual grasses [40]. One possible factor contributing to the decline in California sagebrush density is nitrogen deposition [5,9]. Approximately 45 kg/ha/yr of nitrogen is deposited in coastal sage scrub communities from the Los Angeles Air Basin. As much as 87 kg/ha/yr of nitrogen deposition has been recorded in coastal sage scrub near the urban areas [5,9]. In controlled trials, red brome (Bromus rubens) and rattail fescue overwhelmingly "outcompeted" California sagebrush plants when subjected to elevated levels of nitrogen [6,7,9].

SEASONAL DEVELOPMENT:
California sagebrush initiates new leaf growth with the onset of the 1st significant rainfall. Flower development, branch elongation, and fruit production do not begin for several months after leaf growth begins, with some variation among years and habitats [56,57,63,68,146]. California sagebrush foliage is lush and green, and growth is continuous during rainy periods in winter and spring [42,146,150]. From summer until the onset of winter rains, the large leaves that are produced during the rainy season are dropped, and smaller leaves emerge in late spring and are retained during the dry season [57,63,93,199,203]. Leaf shedding allows California sagebrush to avoid long-term water stress [54,146]. California sagebrush generally shows substantial leaf biomass production within 2 weeks following the end of drought season [170]. Under severe drought conditions, California sagebrush stems become bare and die back [30,93]. California sagebrush shows no measurable aboveground growth for the 2 to 4 driest months of the year [68]. Various flowering times for California sagebrush have been reported; depending upon elevation and geographical location, they include September and October [72], August to December [155,156], November to mid-May [150], year-round [48], and April to October and April to June in Baja California [65,210].

The approximate phenological development of California sagebrush at Mesa La Misin, Baja California, Mexico, is as follows [65]:

Phenological stage Time period
Vegetative growth January to April
Flowering April to June
Fruiting June to August
Dried leaves August to December

FIRE ECOLOGY

SPECIES: Artemisia californica
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: California sagebrush establishes after fire by seed [62,108,109,110,146,200,201,204] and sprouting from the root crown [45,78,125,126,127,128,129,130,200,201]. Vegetative sprouting is the primary postfire regeneration method, occurring in the 1st year after fire [146,204]. Seed dormancy can be broken by light, charate, or a combination of those factors; in part, that response is likely an adaptation to fire [17,108,109,110,146,161].

Fire regimes: There is some disagreement over the historic fire-return interval for coastal sage scrub communities. At one end of the spectrum, Paysen and others [167] estimate the fire return interval to range from <35 to <100 years. At the other end of the spectrum, O'Leary [160] believes the natural fire frequency in coastal sage scrub is near the lower end of a 20- to 40-year cycle common in adjacent chaparral communities. Vogl [194] estimates an average fire-return interval of 20 years for lightning-ignited fire in chaparral adjacent to coastal sage scrub. While the historic fire-return interval may have ranged from ~20 to 100 years, loss of historic range due to development, agriculture, human-caused fires, and vegetation type conversion has altered fire regimes in coastal sage scrub communities [5,40,51].

Anthropogenic influence in coastal sage scrub has altered the fire-return interval to as short as 5 to 10 years or less on some sites [39,160]. In Santa Monica Mountains coastal sage scrub communities, fire data (all fires 0.1 ha) were assessed for the period 1909 to 1977. For the time period covered (68 years), the average interval between fires was 14 years for coastal sage scrub [133]. For the same time period, Hanes [85] suggested the fire return interval in coastal sage scrub communities in the San Gabriel and San Bernardino mountains was 2 to 10 years. The introduction of Mediterranean annual grasses, which often provide continuous ground cover of flash fuels, has severely altered the historic fire regime in coastal sage scrub [66]. Prior to 1960, the historic fire-return interval was 30 years in coastal sage scrub communities in western Riverside County. Over the past 20 years, however, nonnative grass cover has increased and altered the fire-return interval to less than 8 years [40] and likely as short as 2 to 5 years [5,6].

Shortened fire-return intervals may greatly reduce California sagebrush coverage [146]. Computer modeling conducted by Malanson [127] suggests that California sagebrush is eliminated from the shrub canopy within 100 years when fire-return intervals are less than 40 years. Other modeling predicts California sagebrush can handle fire intervals no shorter than 10 years [21]. While decreased fire frequency is of concern, it is likely that California sagebrush can withstand fire-return intervals shorter than every 40 years. California sagebrush is a common understory species in Monterey pine (Pinus radiata) forests at Swanton Pacific Ranch. Using fire scar data, Stephens and others [183] estimate that the mean fire-return interval for the years 1893 to 1976 ranged from 11.4 to 20.1 years. In the Santa Monica Mountains, California sagebrush cover is greatest (32.6% to 50.8%) on sites that burn approximately every 21 to 22 years [125]. For a California sagebrush-eastern Mojave buckwheat community on the Cleveland National Forest, fire records show that stand-replacing fire occurs at approximately 28-year intervals [208].

The following table provides fire return intervals for plant communities and ecosystems where California sagebrush 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)
California chaparral Adenostoma and/or Arctostaphylos spp. 20 to 100 [5,7,40]
coastal sagebrush Artemisia californica <35 to <100
California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100 [167]
California steppe Festuca-Danthonia spp. <35
pinyon-juniper Pinus-Juniperus spp. <35 [167]
California oakwoods Quercus spp. <35 [11]
coast live oak Quercus agrifolia 2-75 [77]


POSTFIRE REGENERATION STRATEGY [184]:
Small shrub, adventitious bud/root crown
Ground residual colonizer (on-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

FIRE EFFECTS

SPECIES: Artemisia californica
 

 

Postfire sprouting California sagebrush on Badger Hill, California State University, San Bernardino. Photo Courtesy of Michael Loik

IMMEDIATE FIRE EFFECT ON PLANT:
Low-severity fire top-kills California sagebrush [43,118,215]; high-severity fire causes mortality [111,200,216,217]. While the effect of medium-severity fires is not discussed in the literature, Zedler [215] rates California sagebrush mortality from fire as "intermediate" to "high". Shallowly buried seeds are often killed by fire [129,200]; deeply buried seeds survive fire [198].

California sagebrush has a high ratio of dead:live material, making it highly flammable [25]. Shrub species with a high proportion of ether extracts, above 8%, are also considered highly flammable. California sagebrush has a 15.6% level of ether extracts [74,148], characterizing it as "explosively flammable" [148,204].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No additional information is available on this topic.

PLANT RESPONSE TO FIRE:
California sagebrush establishes after fire by seed [62,78,108,109,110,146,177,200,201,204] and sprouting from the root crown [45,78,78,125,126,127,128,129,130,200,201]. Root crown sprouting is the primary postfire regeneration method, occurring in the 1st year after fire. California sagebrush seedling emergence after fire is considered "variable" and generally "low" [146,204]. Seed production does not usually take place on burn sites until after the sprouts have flowered and set seed, in postfire year 2 at the earliest [108,204]. However, seedling establishment on burned sites in the 1st postfire year may occur from soil-stored seed [108,198] not killed by high-severity fire [129,200] or seeds dispersed onto the burned site by wind [54,56,57,141,146,150] or animal activity [56,57]. Sprouting California sagebrush plants flower vigorously the 1st few years after burning [160,214]. Sprouting tends to be less in burn areas composed of dense vegetation, where California sagebrush plants are older, or where fire severity was high [146]. Westman and others [200] note that coastal California sagebrush plants tend to recover after fire more quickly than inland plants. California sagebrush coastal populations experience greater precipitation and more temperate conditions than do inland plants, creating more suitable conditions for recovery following fire [40,84,93].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Increased postfire production Three years following a prescription fire in a Morro manzanita (Arctostaphylos morroensis) community, California sagebrush cover and frequency substantially increased. The burn, located in Montaa de Oro State Park, was conducted on 4 November 1998. Air temperature during the fire was 61 F (16C), and relative humidity ranged from 50% to 60%. Flame lengths as great as 50 feet (15 m) were observed. In places, litter was still smoldering 24 hours following ignition. Prior to the fire, California sagebrush cover and frequency were 0.4% and 8.0%, respectively. In postfire year 3, cover and frequency increased to 7.0% and 61.0%, respectively. As a result of the burn, pure stands of Morro manzanita were transformed to mixed stands of native California sagebrush, coyote bush, and orange bush monkeyflower (Diplacus aurantiacus), and nonnative hottentot fig (Carpobrotus edulis), perennial veldtgrass (Ehrharta calycina), and narrow-leaved iceplant (Conicosia pugioniformis) [162].

On Otay Mountain, in southwest San Diego County, 2 "high-severity" wildfires during the summers of 1979 and 1980 caused high mortality of California sagebrush. The fire in August 1979 caused >95% mortality of California sagebrush plants. During the summer of 1980, an arsonist set fire to a portion of the site that burned in 1979. Where fire burned in both 1979 and 1980, the mortality rate of California sagebrush was >96%. Prior to the 1979 and 1980 fires, the density (no./msx) of mature California sagebrush shrubs was 0.220.06 on sites that burned only in 1979 and 0.260.06 on sites burned in 1979 and 1980. While no data exist on California sagebrush seedlings prior to the 1979 fire, seedling densities after the 1979 and 1980 fire where 2.620.30/m and 0.390.12/m, respectively. When combining seedlings and mature shrubs that survived fire, the relative abundance of California sagebrush in May 1981 was 9.20 times greater on sites burned only in 1979 and 0.78 times greater on sites burned in both 1979 and 1980 compared to abundance of mature shrubs before fire at both sites. Since the mortality rate of mature shrubs was so high, nearly all California sagebrush observed in 1981 were seedlings and not sprouts [216,217].

Postfire sprout/seedling regeneration: Sprouting of California sagebrush in postfire year 1 is common, but seedling establishment generally does not occur until at least postfire year 2. The Agoura-Kanan Fire, located in the Santa Monica Mountains, burned on 23 October 1978. The 2 study sites investigated, Encinal Canyon and Kanan-Dume, had been fire free for 22 and 11 years, respectively. Both sites are located on the coastal side of the range. California sagebrush sprouting and seedling recruitment were assessed in May 1979 (postfire month 7) and May 1980 (postfire month 19). There were no seedlings in postfire year 1, but sprouts flowered and set seed in abundance in year 1, facilitating a large flush of seedlings in postfire year 2 in Encinal Canyon. The following table reflects California sagebrush sprouts and seedlings as a percent of total community density at both sites in postfire years 1 and 2 [114].

  Kanan-Dume Encinal Canyon
May 1979 May 1980 May 1979 May 1980
% sprouts 13 19 1 5
% seedlings 0 1 0 30

Total seedling establishment on 6 burned sites far outpaced California sagebrush sprouts. Two years following a late October 1978 fire that burned 20,000 acres (10,000 ha) on the coastal side of the Santa Monica Mountains, Malanson and O'Leary [128] sampled the number of California sagebrush sprouts and seedlings on 6 burn sites. Prior to the 1978 fire, the sites had not burned for 22 years. Site 1 is on south-facing sandstone; site 2 is on north-facing sandstone; sites 2 and 4 are on east-facing sandstone; and sites 5 and 6 are on west-facing shale. Sites 1 through 4 burned in mid-afternoon during a period of low humidity (10-15%). Sites 5 and 6 burned at 2 a.m., with a relative humidity from of 40% to 45% [128].

Site 1 Site 2 Site 3 Site 4 Site 5 Site 6
Sprout (no. /ha) 3,437 312 312 312 781 625
Seedlings (no. /ha) 7,500 156 312 56,719 469 6,250

Malanson and O'Leary 1985 [129] returned to the burn site at postfire year 4.5 to measure the canopy size of California sagebrush on the 6 sites. California sagebrush canopy size was assessed on high intensity (6,100 kcal/m) and low intensity (5,600 kcal/m) burn sites, on andesite and sandstone substrates, and on south-, north-, and east-facing slopes. California sagebrush canopy was greatest on low-intensity burns, on andesite substrates, and south-facing slopes. Of the 5 species measured (California brittle bush, black sage, coastal buckwheat (Eriogonum cinereum), and San Luis purple sage), California sagebrush had the smallest total canopy size of all species. California sagebrush canopy coverage was [129]:

Site characteristics Canopy size (m)
High intensity burn 779 (6,327*)
Low intensity burn 962 (9,327)
Andesite substrate 985 (5,623)
Sandstone substrate 419 (5,984)
South-facing slope 985 (6,105)
North-facing slope 107 (2,206)
East-facing slope 597 (5,774)
*total canopy size of all 5 species combined

Postfire comparisons in coastal and interior sage scrub communities: Over a 10-day period in late October and early November 1993, over 200,000 acres (80,000 ha) of chaparral and coastal sage scrub were burned by wildfire. The severity of the fire ranged from low to high and affected areas that had not burned from 3 to 85 years ago. On coastal sage scrub burned sites, peak vegetative recovery occurred in late spring. During late spring, the ratio of California sagebrush seedlings to prefire shrubs was approximately 15:1, and the ratio of California sagebrush sprouts:prefire shrubs was approximately 0.3:1. The lower ratio of sprouts to prefire shrubs was due in large part to complete fire mortality of many California sagebrush shrubs. Measurements were also taken of the basal diameter and height of both nonsprouting (dead) and sprouting (live) postfire California sagebrush skeletons. Not surprisingly, the height of sprouting California sagebrush shrubs was significantly (p<0.001) greater than the height of nonsprouting shrubs. The basal diameter of sprouting shrubs was significantly (p<0.001) less than nonsprouting shrubs. This suggests that juvenile shrubs are better suited to survive and sprout following fire than older shrubs. The numbers in parentheses in the following table indicate the number of California sagebrush shrubs sampled [111].

Shrub status

Basal diameter
(mmsx)

Height
(cmsx)

Nonsprouting 321 (952) 191 (951)
Sprouting 111 (376) 322 (376)

Following the 1993 fires, the percentages of sprouting California sagebrush plants and the seedling:parent ratio were evaluated in postfire year 1 on coastal sage scrub and interior sage scrub communities. The number of parents was based on the census of shrub skeletons, and seedling ratios were evaluated only for sites with more than 5,000 seedlings/ha. Of the 11 shrub skeletons present in coastal sage scrub and 19 in interior sage scrub, 18%8% (xs) and 20%6% had sprouted following fire, respectively. Of the 2 parent plants found in coastal sage scrub and 23 in interior sage scrub, 150 (xs) and 3310 seedlings/plant were measured, respectively [113].

The total California sagebrush seedling recruitment for all 5 years (no. of seedlings/has x) in chaparral communities following the 1993 fire was 11,6004,800 and 31,50010,600 in sage scrub communities. Approximately 83% of all seedlings recruited established during postfire years 1 and 2 [113].

Postfire diversity: One year after the 1993 fires, species densities (no. of species/0.1 hasx) in coastal sage scrub and interior sage scrub communities codominated by California sagebrush were 35.42.1 (sx) and 58.52.4, respectively. At the end of postfire year 5, species density in coastal sage scrub and interior sage scrub communities averaged 72.31.9 and 94.72.4, respectively. Further, by the end of postfire year 5, coastal sage scrub communities were significantly (p<0.05) closer to their prefire plant density than in postfire year 1. While interior sage scrub community densities were more similar in postfire year 5 to prefire levels than during postfire year 1, the change was not significant [112].

FIRE MANAGEMENT CONSIDERATIONS:
Avian community dynamics: Several studies have found that bird species preferentially chose unburned coastal sage scrub communities over recently burned sites. Stanton [182] studied bird populations in a burned and unburned coastal sage scrub community 2 years following prescribed burning in Los Angeles County. While many bird species were found at both sites, more species and individuals selected the unburned site, which offered better foraging opportunities. On the unburned site, the bird population remained relatively steady throughout the year. The population on the burned site fluctuated, with greatest use in the spring [182].

Late-successional California sagebrush communities provide critical habitat for the federally threatened [192] California gnatcatcher [21,132,212]. California gnatcatcher nesting sites were surveyed at coastal sites in Riverside and San Diego counties. Burned and adjacent unburned plots, which had not burned for 22 years, were sampled at postfire year 3. California gnatcatcher nests were found at all unburned sites (n≥5). California sagebrush cover on unburned sites ranged from 4.2% to 78.4%. No nests were found at the burned sites (n=7) in postfire year 3, and California sagebrush cover ranged from a trace to 11.8%. California gnatcatcher nesting site requirements include a shrub cover of at least 50% and a minimum average shrub height of 3 feet (1 m); neither of these requirements was met on the 7 recently burned sites [21]. Beyers and Pea [22] and Mayer and Wirtz [132,212] conducted similar studies within burned and unburned sites. In both studies, California gnatcatcher highly preferred unburned sites.

For 1 year following a wildfire in Box Canyon, Los Angeles County, bird populations were counted and compared in burned and unburned areas. Bird censuses were taken 20 times during the year following the fire. Of the 72 species seen at both sites, slightly but not significantly more birds were concentrated on the unburned site [151].

Fire suppression: In coastal sage scrub, fire suppression has generally been successful in summer months devoid of extremely high temperatures accompanied by Santa Ana winds in fall. Where fire suppression has been successful year-round, delayed burning has resulted in abnormally large fires when the abovementioned weather conditions occur [127]. Coastal sage scrub is generally closer to developed areas than chaparral and is thus susceptible to accidentally and arsonist-set fire, thus increasing fire suppression activities. The long-term effect of fire exclusion on California sagebrush and other coastal sage scrub species in unknown. Zedler [214] stated that increased fire frequency in some coastal sage scrub communities is likely not natural, and fire suppression may be necessary to restore the communities to presettlement conditions. Conversely, some researchers suggest that fire exclusion facilitates the invasion of coastal sage scrub species into adjacent communities. Epling and Lewis [58] state that California sagebrush and other coastal sage scrub species are successfully invading chaparral communities dominated by sprouting species following abnormally hot, stand-replacement fires resulting from fuel buildup facilitated by suppression measures. Research conducted by Thomas [187] found that California sagebrush is invading adjacent valley oak (Quercus lobata) woodlands in the Santa Monica Mountains National Recreation Area.

Fuel loading: California sagebrush dominated communities biomass varies from 3 to less than 15 tons/acre, and approximately 70% to 85% of the biomass is consumed by hot fire [75,159]. In a California sagebrush-common deerweed community, total aboveground fuel load was 13.2 tons/acre. Aboveground live and dead biomass was 3.7 and 2.8 tons/acre, respectively, and duff and litter biomass was 6.7 tons/acre [159].

Prescription burning: Prescription burning of California sagebrush is not advised for several reasons. Since fire can deplete seed banks and kill adult plants, regular burning may reduce California sagebrush [127]. Further, prescription burning may favor further encroachment of Mediterranean grasses (red brome, ripgut brome, wild oats (Avena spp.), and annual sixweeks grasses (Vulpia spp.)) into California sage scrub communities [5]. If prescription burning is undertaken in coastal sage scrub, spring is the favorable season since less heat is generated by fires [72]. Guidelines for developing a fire prescription management plan in coastal sage scrub are available in the literature [74,75].

MANAGEMENT CONSIDERATIONS

SPECIES: Artemisia californica
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
The pungent aroma and bitter taste of California sagebrush is unpalatable to livestock and wildlife [176]. DeSimone and Zedler [54] suggest that terpenes contained in the leaves of California sagebrush protect the plant from herbivory. Columbian black-tailed deer browse California sagebrush "limitedly" during the growing season [176]. Historically, feral domestic sheep and goats browsed California sagebrush on south-facing slopes of the Channel Islands [32,141]. In the Los Padres National Forest, domestic goats are used to manage brush regrowth on fuelbreaks. Where California sagebrush occurs, domestic goats generally browse on the shrub year-round [76].

During the dry season in northern Baja California, domestic goats browse California sagebrush and bastardsage (Eriogonum wrightii). Domestic goats feed on California sagebrush during leaf initiation, vegetative growth, flowering, fruiting, and dried leaf stages. Feeding mostly occurs during the dried leaf stage [65].

Small mammals/birds: The California gnatcatcher preferentially forages in California sagebrush and eastern Mojave buckwheat stands, particularly because the shrubs support a high number and variety of arthropods [13,21,146]. Arthropods are more abundant on California sagebrush than on eastern Mojave buckwheat [13].

Dusky-footed and desert woodrats have a high food preference for California sagebrush [136]. Near Irvine, 7 rodent species (4 cricetids and 3 heteromyids) fed regularly on California sagebrush from early winter until early spring [137].

Palatability/nutritional value: Due to the pungent aroma and bitter taste of California sagebrush, palatability of the plant is low [176]. While little browsed, California sagebrush protein content is adequate for livestock production [65].

The crude protein content of California sagebrush at Mesa La Mision, Baja California, during 5 phenological stages is presented below [65]:

Phenological stage Crude protein content (%)
Leaf initiation 14.3
Vegetative growth 10.4
Flowering 14.1
Fruiting 6.9
Dry leaves 6.0

The mean (8 samples/month for 24 months) annual nutrient content of California sagebrush in the Santa Monica Mountains is presented below. Plants were from a site 490 feet (150 m) above sea level, 2 miles (3 km) from the ocean, and on a 10% to 15% slope [69,71].

N (%) P (%) K (%) Ca (%) Mg (%)
1.88 0.225 1.42 0.94 0.267

Cover value: As the tallest subshrub in the coastal sage scrub community, California sagebrush is an important cover species for a variety of small birds, mammals, amphibians, and arthropods [30,33,81,104,123,166,190,193]. In a review, Wirtz [211] lists common mammals, amphibians, arthropods, and birds associated with California sagebrush.

California sagebrush provides preferred habitat for Bell's sage sparrow, a bird species endemic to shrubby habitats and state-listed as a Species of Special Concern [3]. At the Motte Rimrock Reserve, successful Bell's sage sparrow nesting sites are associated with areas high in California sagebrush and brittle bush cover [145].

The California gnatcatcher is found almost exclusively in California sagebrush-dominated communities [2,21,146,174]. In southern California, California gnatcatcher nests are most frequently found in California sagebrush stands, followed in order by white sage, black sage, chamise, and prickly-pear (Opuntia ssp.) [13,21].

VALUE FOR REHABILITATION OF DISTURBED SITES:
California sagebrush seeds and transplants have been used successfully in the rehabilitation of disturbed sites [27,33,40,46,67,91,96,97,106,165,197]. Further, California sagebrush is described as a shrub that has erosion control potential [34,102,147].

OTHER USES:
California sagebrush is used as an ornamental [31] and has an extensive history as a plant of importance for Native Americans [146], miners [48], and Spanish [48] and Mexican settlers [79]. The pollen of California sagebrush is extracted and used as a desensitizing agent for the relief of hay fever [79].

California sagebrush is considered 1 of the most important medicinal plants by Cahuilla Native Americans in southern California. It was used to induce menstruation, provide a comfortable child-birth experience, and promote rapid postnatal recovery. Beginning with the onset of menstruation, young girls were given a tea made from boiled California sagebrush. From this point forward, California sagebrush tea was drunk just prior to each menstrual period for the duration of their reproductive life. The drink was given to newborn babies 1 day after birth to clean out their system [19,146]. The leaves of California sagebrush were also used by Cahuilla Native Americans to relieve colds, chewed fresh or dried, and were smoked after mixing with tobacco and other leaves. California sagebrush was also used in Cahuilla sweathouses for various cures [19,146].

The Chumash Native Americans of southern California had many uses for California sagebrush. The branches were used as firesticks, arrow foreshafts, and brush windbreaks and enclosures. California sagebrush was processed into a poultice and applied directly to the forehead to relieve headaches. The Chumash would soak California sagebrush leaves in water and bath in it or use it as a ritualistic purification sprinkling agent, particularly related to death. Bundles of branches were also erected along paths to shrines [189].

OTHER MANAGEMENT CONSIDERATIONS:
Fertilization: Nitrogen fertilization of California sagebrush seedlings caused a significant (p<0.01) increase in shoot biomass. Soil was amended with nitrogen concentrations of 2 g/g, 20 g/g, 40 g/g, and 80 g/g. Following 3 months of growth, California seedling shoots at the 4 treatments had a biomass (oven dried) of 0.07 g, 0.67 g, 0.96 g, and 1.18 g, respectively [163].

Browsing: Heavy browsing of California sagebrush by feral domestic sheep on south-facing slopes of Santa Cruz Island has caused the complete destruction of the plant community [32]. California sagebrush grows best in areas protected from domestic goats [142].

Growth regulators: A reduction in California sagebrush growth over a 3-year period occurred following the application of several growth regulator chemicals (PP333 or EL500). Growth regulators were applied from 1981 to 1983 to evaluate there use in reducing fuel to prevent wildfires. In 4 field trials, California sagebrush height was significantly (p<0.05) reduced by growth regulators [95].

Herbicides: Glyphosate concentrations from 1.0% to 2.0% resulted in the death of 90% to 99% of California sagebrush shrubs 1 year following treatment in southern California [105]. California sagebrush plants can be controlled and/or killed with the herbicides 2,4-D [164].

Loss of range: Estimates of California sagebrush-dominated habitat vary. Several authors report that California sagebrush-dominated coastal sage scrub has lost 66% to 90% of its original vegetation to development, agriculture, vegetation type conversion, and changing fire regimes [5,40,51]. Conversely, Taylor [186], using U.S. Forest Service Vegetation Type Maps, assessed the loss of California sagebrush dominated and codominated habitat in southern California from 1931 to 1995. The researcher estimates that 64% of California sagebrush communities remained undeveloped in 1995, while 59% of California sagebrush-eastern Mojave buckwheat-chamise plots remained undeveloped in 1995 [186].

Nonnative species: The invasion of coastal sage scrub communities by Mediterranean annual grasses may be increasing [5]. The annual grasses soft chess (Bromus hordeaceus), red brome, ripgut brome, slender oat (Avena barbata), and wild oat (A. fatua) may depress growth of young California sagebrush plants. The question of possible growth interference was evaluated at the Santa Margarita Ecological Reserve near San Diego. In plots without annual grasses, aboveground biomass of 4-month-old California sagebrush seedlings was significantly (p<0.05) greater than in plots with annual grasses. At 16 months of age, California sagebrush plant biomass remained significantly less in plots with annual grasses. California sagebrush plants may have had less biomass when cooccurring with annual grasses due to depletion in available water and sunlight [57]. However, by the end of the 2nd growing season, the biomass of surviving California sagebrush plants was similar to that of plants in plots without annual grasses [56].

Prescription fire is discouraged as a method of controlling nonnative species in California sagebrush communities. Alternative methods are currently being researched. From 1999 to 2002, Allen [5] found that control of Mediterranean grasses was possible via domestic sheep grazing and grass-specific herbicide application.

Plant facilitation: California sagebrush has a positive influence on the establishment of blue oak (Quercus douglasii) seedlings. In Carmel Valley, Monterey County, blue oak seedlings were significantly (p<0.001) associated with California sagebrush and San Luis purple sage. While  combined cover of California sagebrush and San Luis purple sage was 37% at the study site, 91% of all blue oak seedlings were located under the canopies of the 2 shrubs. Shade provided by the shrub canopy promotes blue oak seedling establishment [35].

In the San Ynez Valley and San Ynez Mountains, California sagebrush and San Luis purple sage have a positive influence on coast live oak (Q. agrifolia) seedlings. At the 2 study sites, coast live oak seedlings were significantly (p<0.001) associated with California sagebrush and San Luis purple sage. Combined cover of the 2 shrubs at the 2 sites was 36% and 27% on San Ynez Valley and San Ynez Mountain plots, respectively, However, 80% and 68% of all coast live oak seedlings on San Ynez Valley and San Ynez Mountain plots, respectively, were found under the canopies of California sagebrush and San Luis purple sage. At these 2 sites, shade provided by canopy cover was not the only contributing factor for coast live oak establishment and survivorship. Significantly (p<0.05) fewer coast live oak seedlings growing under the shrub canopies were browsed by livestock, mule deer, and California pocket gophers compared to seedlings growing in open grassland or under the canopy of adult coast live oak trees. The percentages of seedlings browsed under the canopies of California sagebrush and San Luis purple sage were 7.714.0 (xs) and 8.714.7, respectively, in open grasslands and 52.932.2 and 50.035.9, respectively, under coast live oak [36]. A similar study by Callaway and Davis [38], conducted in Gaviota State Park, evaluates the importance of shrubs (including California sagebrush) in the recruitment of coast live oak seedlings.

Soil amendments: On disturbed land on the Santa Margarita Ecological Reserve, soil was amended with a slowly decaying amendment (pine bark) and a rapidly decaying amendment (oat straw) prior to the planting of 2-inch (5 cm) California sagebrush seedlings. The survival rate of California sagebrush seedlings 28 months after planting was ~25% in the pine bark, ~16% in oat straw, and ~15% in the control. Plant volume 28 months after planting in pine bark, oat straw, and control treatments was ~275,000 cm, ~125,000 cm, and 100,000 cm, respectively [218].

Artemisia californica: REFERENCES


1. Agee, James K.; Biswell, Harold H. 1978. The fire management plan for Pinnacles National Monument. In: Linn, Robert M., ed. Proceedings, 1st conference on scientific research in the National Parks: Vol. 2; 1976 November 9-12; New Orleans, LA. NPS Transactions and Proceedings No. 5. Washington, DC: U.S. Department of the Interior, National Park Service: 1231-1238. [14368]
2. Akcakaya, H. Resit; Atwood, Jonathan L. 1997. A habitat-based metapopulation model of the California gnatcatcher. Conservation Biology. 11(2): 422-434. [63457]
3. Akcakaya, H. Resit; Franklin, Janet; Syphard, Alexandra D.; Stephenson, John R. 2005. Viability of Bell's sage sparrow (Amphispiza belli ssp. belli): altered fire regimes. Ecological Applications. 15(2): 521-531. [54517]
4. Allen, Barbara H.; Holzman, Barbara A.; Evett, Rand R. 1991. A classification system for California's hardwood rangelands. Hilgardia. 59(2): 1-45. [17371]
5. Allen, Edith B. 2004. Restoration of Artemisia shrublands invaded by exotic annual Bromus: a comparison between southern California and the Intermountain Region. In: Hild, Ann L.; Shaw, Nancy L.; Meyer, Susan E.; Booth, D. Terrance; McArthur, E. Durant, compilers. Seed and soil dynamics in shrubland ecosystems: proceedings; 2002 August 12-16; Laramie, WY. Proceedings RMRS-P-31. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 9-17. [48994]
6. Allen, Edith B.; Eliason, Scott A.; Marquez, Viviane J.; Schultz, Gillian P.; Storms, Nancy K.; Stylinski, Cathlyn Davis; Zink, Thomas A.; Allen, Michael F. 2000. What are the limits to restoration of coastal sage scrub in southern California? In: Keeley, J. E.; Keeley, M. B.; Fotheringham, C. J., eds. 2nd interface between ecology and land development in California. USGS Open-File Report 00-62. Sacramento, CA: U.S. Department of the Interior, Geological Survey, Western Ecological Research Center: 253-262. [47657]
7. Allen, Edith B.; Padgett, Pamela E.; Bytnerowicz, Andrzej; Minnich, Richard. 1998. Nitrogen deposition effects on coastal sage vegetation of southern California. In: Bytnerowicz, Andrzej; Arbaugh, Michael J.; Schilling, Susan L., tech. coords. Proceedings of the international symposium on air pollution and climate change effects on forest ecosystems; 1996 February 5-9; Riverside, CA. Gen. Tech. Rep. PSW-GTR-166. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 131-139. [64971]
8. Allen, Edith B.; Sirulnik, Abby G.; Egerton-Warburton, Louise; Kee, Sheila N.; Bytnerowicz, Andrzej; Padgett, Pamela E.; Temple, Patrick J.; Fenn, Mark E.; Poth, Mark A.; Meixner, Thomas. 2005. Air pollution and vegetation change in southern California coastal sage scrub: a comparison with chaparral and coniferous forest. In: Kus, Barbara E.; Beyers, Jan L., tech. coords. Planning for biodiversity: Bringing research and management together: Proceedings of a symposium for the south coast ecoregion; 2000 February 29 - March 2; Pomona, CA. Gen. Tech. Rep. PSW-GTR-195. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 79-95. [64505]
9. Allen, Michael F.; Egerton-Warburton, Louise; Treseder, Kathleen K.; Cario, Cara; Lindahl, Amy; Lansing, Jennifer; Querejeta, Jose Ignacio; Karen, Ola; Harney, Sharon; Zink Thomas. 2005. Biodiversity of mycorrhizal fungi in southern California. In: Kus, Barbara E.; Beyers, Jan L., tech. coords. Planning for biodiversity: Bringing research and management together: Proceedings of a symposium for the south coast ecoregion; 2000 February 29 - March 2; Pomona, CA. Gen. Tech. Rep. PSW-GTR-195. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 43-56. [64502]
10. Allen-Diaz, Barbara H.; Holzman, Barbara A. 1991. Blue oak communities in California. Madrono. 38(2): 80-95. [15424]
11. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]
12. Arno, Stephen F.; Wilson, Andrew E. 1986. Dating past fires in curlleaf mountain-mahogany communities. Journal of Range Management. 39(3): 241-243. [350]
13. Atwood, Jonathan L. 1993. California gnatcatchers and coastal sage scrub: the biological basis for endangered species listing. In: Keeley, Jon E., ed. Interface between ecology and land development in California: Proceedings of the symposium; 1992 May 1-2; Los Angeles, CA. Los Angeles, CA: The Southern California Academy of Sciences: 149-169. [21704]
14. Axelrod, Daniel I. 1978. The origin of coastal sage vegetation, Alta and Baja California. American Journal of Botany. 65(10): 1117-1131. [5563]
15. Baker, Herbert G. 1972. Seed weight in relation to environmental conditions in California. Ecology. 53(6): 997-1010. [63466]
16. Bartolome, James W. 1989. Local temporal and spatial structure. In: Huenneke, L. F.; Mooney, H., eds. Grassland structure and function: California annual grassland. Dordrecht, The Netherlands: Kluwer Academic Publishers: 73-80. [28348]
17. Baskin, Carol C.; Baskin, Jerry M. 2001. Seeds: ecology, biogeography, and evolution of dormancy and germination. San Diego, CA: Academic Press. 666 p. [60775]
18. Bauer, Harry L. 1936. Moisture relations in the chaparral of the Santa Monica Mountains, California. Ecological Monograph. 6(3): 409-454. [10528]
19. Bean, Lowell John; Saubel, Katherine Siva. 1972. Telmalpakh: Chauilla Indian knowledge and usage of plants. Banning, CA: Malki Museum. 225 p. [35898]
20. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
21. Beyers, Jan L., Wirtz, William O., II. 1997. Vegetative characteristics of coastal sage scrub sites used by California gnatcatchers: implications for management in a fire-prone ecosystem. In: Greenlee, Jason M., ed. Proceedings, 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 81-89. [28126]
22. Beyers, Jan L.; Pena, Ginger C. 1995. Characteristics of coastal sage scrub in relation to fire history and use by California gnatcatchers. In: Weise, David R.; Martin, Robert E., technical coordinators. The Biswell symposium: fire issues and solutions in urban interface and wildland ecosystems; 1995 February 15-17; Walnut Creek, CA. Gen. Tech. Rep. PSW-GTR-158. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 153-154. [26940]
23. Beyers, Jan L.; Stewart, Theresa A.; Sharp, Cary. 1995. A postfire seeding experiment at the San Diego Wild Animal Park. In: Keeley, Jon F.; Scott, Tom, eds. Brushfires in California: ecology and resource management: Proceedings; 1994 May 6-7; Irvine, CA. Fairfield, WA: International Association of Wildland Fire: 199-204. [43343]
24. Bjorndalen, Jorn Erik. 1978. The chaparral vegetation of Santa Cruz Island, California. Norwegian Journal of Botany. 25: 255-269. [7851]
25. Bolsinger, Charles L. 1989. Shrubs of California's chaparral, timberland, and woodland: area, ownership, and stand characteristics. Res. Bull. PNW-RB-160. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Experiment Station. 50 p. [7426]
26. Bolton, Robert B., Jr.; Vogl, Richard J. 1969. Ecological requirements of Pseudotsuga macrocarpa in the Santa Ana Mountains, California. Journal of Forestry. 67: 112-116. [10807]
27. Bowcutt, Frederica. 1990. Restoring coastal sage scrub at San Onofre State Beach (California). Restoration & Management Notes. 8(2): 120. [13679]
28. Boyd, Steve. 1999. Vascular flora of the Liebre Mountains, western Transverse Ranges, California. Aliso. 18(2): 93-139. [40639]
29. Bradbury, David E. 1978. The evolution and persistence of a local sage/chamise community pattern in southern California. Yearbook of the Association of Pacific Coast Geographers. 40: 39-56. [12269]
30. Brown, David E., ed. 1982. Biotic communities of the American Southwest--United States and Mexico. Desert Plants: Special Issue. Tucson, AZ: University of Arizona Press. 4(1-4): 1-342. [62041]
31. Browse, Philip McMillan. 1987. Artemisia californica 'Canyon Gray'. Pacific Horticulture. 48(3): 56. [63321]
32. Brumbaugh, Robert W.; Leishman, Norman J. 1982. Vegetation change on Santa Cruz Island, California: the effect of feral animals. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 589. [6064]
33. Burger, J. C.; Redak, R. A.; Allen, E. B.; Rotenberry, J. T.; Allen, M. F. 2003. Restoring arthropod communities in coastal sage scrub. Conservation Biology. 17(2): 460-467. [63322]
34. Burkhart, Brad. 1988. A nurseryman's view of revegetation. In: Rieger, John P.; Williams, Bradford K., eds. Proceedings of the second native plant revegetation symposium; 1987 April 15-18; San Diego, CA. Madison, WI: University of Wisconsin Arboretum, Society for Ecological Restoration & Management: 17-21. [4089]
35. Callaway, Ragan M. 1992. Effect of shrubs on recruitment of Quercus douglasii and Quercus lobata in California. Ecology. 73(6): 2118-2128. [18431]
36. Callaway, Ragan M.; D'Antonio, Carla M. 1991. Shrub facilitation of coast live oak establishment in central California. Madrono. 38(3): 158-169. [17102]
37. Callaway, Ragan M.; Davis, Frank W. 1993. Vegetation dynamics, fire, and the physical environment in coastal central California. Ecology. 74(5): 1567-1578. [21675]
38. Callaway, Ragan M.; Davis, Frank W. 1998. Recruitment of Quercus agrifolia in central California: the importance of shrub-dominated patches. Journal of Vegetation Science. 9(5): 647-656. [41369]
39. Carrington, M. E.; Keeley, J. E. 1999. Comparison of post-fire seedling establishment between scrub communities in mediterranean and non-mediterranean climate ecosystems. Journal of Ecology. 87(6): 1025-1036. [37577]
40. Cione, Nancy K.; Padgett, Pamela E.; Allen, Edith B. 2002. Restoration of a native shrubland impacted by exotic grasses, frequent fire, and nitrogen deposition in southern California. Restoration Ecology. 10(2): 376-384. [43206]
41. Clark, Ronilee A.; Halvorson, William L.; Sawdo, Andell A.; Danielsen, Karen C. 1990. Plant communities of Santa Rosa Island, Channel Islands National Park. Tech. Rep. No. 42. Davis, CA: University of California, Institute of Ecology, Cooperative National Park Resources Studies Unit. 93 p. [18246]
42. Conrad, C. Eugene; Roby, George A.; Hunter, Serena C. 1986. Chaparral and associated ecosystems management: a 5-year research and development program. Gen. Tech. Rep. PSW-91. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 15 p. [4885]
43. Cook, Sherburne F., Jr. 1959. The effects of fire on a population of small rodents. Ecology. 40(1): 102-108. [230]
44. Cooper, William Skinner. 1922. The broad-sclerophyll vegetation of California: an ecological study of the chaparral and its related communities. Publ. No. 319. Washington, DC: The Carnegie Institution of Washington. 145 p. [6716]
45. Cox, George W. 1986. Mima mounds as an indicator of the presettlement grassland-chaparral boundary in San Diego County, California. The American Midland Naturalist. 116(1): 64-77. [27432]
46. D'Antonio, Carla M.; Howald, Ann M. 1990. Evaluating the effectiveness of hydroseed mixes, topsoil conservation and other revegetation techniques: a case study in Santa Barbara Co., California. In: Hughes, H. Glenn; Bonnicksen, Thomas M., eds. Restoration `89: the new management challenge: Proceedings, 1st annual meeting of the Society for Ecological Restoration; 1989 January 16-20; Oakland, CA. Madison, WI: The University of Wisconsin Arboretum, Society for Ecological Restoration: 338-348. [14710]
47. Dagit, Rosi. 2002. Post-fire monitoring of coast live oaks (Quercus agrifolia) burned in the 1993 Old Topanga Fire. In: Standiford, Richard B.; McCreary, Douglas; Purcell, Kathryn L., technical coordinators. Proceedings of the 5th symposium on oak woodlands: oaks in California's changing landscape; 2001 October 22-25; San Diego, CA. Gen. Tech. Rep. PSW-GTR-184. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 243-249. [42321]
48. Dale, Nancy. 1986. Flowering plants: The Santa Monica Mountains, coastal and chaparral regions of southern California. Santa Barbara, CA: Capra Press. 239 p. In cooperation with: The California Native Plant Society. [7605]
49. David Magney Environmental Consulting. 2002. Natural vegetation of the Ventura River: Project No. 02-0111, [Online]. In: Baseline conditions draft report (F3) milestone. Appendix D - Environmental impact report. In: Matilija Dam Ecosystem Restoration Feasibility Study. Ventura County Watershed Protection District (Producer). Available: http://www.matilijadam.org/f3/d-alt2.pdf [2005, June 16]. [53567]
50. Davis, Frank W.; Hickson, Diana E.; Odion, Dennis C. 1988. Composition of maritime chaparral related to fire history and soil, Burton Mesa, Santa Barbara County, California. Madrono. 35(3): 169-195. [6162]
51. Davis, Frank W.; Stine, Peter A.; Stoms, David M. 1994. Distribution and conservation status of coastal sage scrub in southwestern California. Journal of Vegetation Science. 5: 743-756. [25944]
52. DeBano, Leonard F. 1999. Chaparral shrublands in the southwestern United States. In: Ffolliott, Peter F.; Ortega-Rubio, Alfredo, eds. Ecology and management of forests, woodlands, and shrublands in the dryland regions of the United States and Mexico: perspectives for the 21st century. Co-edition No. 1. Tucson, AZ: The University of Arizona; La Paz, Mexico: Centro de Investigaciones Biologicas del Noroeste, SC; Flagstaff, AZ: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 83-94. [37047]
53. Desimone, Sandra A.; Burk, Jack H. 1992. Local variation in floristics and distributional factors in Californian coastal sage scrub. Madrono. 39(3): 170-188. [19079]
54. DeSimone, Sandra A.; Zedler, Paul H. 1999. Shrub seedling recruitment in unburned Californian coastal sage scrub and adjacent grassland. Ecology. 80(6): 2018-2032. [30403]
55. DeSimone, Sandra A.; Zedler, Paul H. 2001. Do shrub colonizers of southern Californian grassland fit generalities for other woody colonizers? Ecological Applications. 11(4): 1101-1111. [63349]
56. Eliason, Scott A; Allen, Edith B. 1997. Exotic grass competition in suppressing native shrubland re-establishment. Restoration Ecology. 5(3): 245-255. [27445]
57. Eliason, Scott Andrew. 1995. Competition between Artemisia californica and Mediterranean annual grasses. San Diego, CA: San Diego State University. 64 p. Thesis. [64695]
58. Epling, Carl; Lewis, Harlan. 1942. The centers of distribution of the chaparral and coastal sage associations. The American Midland Naturalist. 27: 445-462. [9793]
59. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
60. Flora of North America Association. 2007. Flora of North America: The flora, [Online]. Flora of North America Association (Producer). Available: http://www.fna.org/FNA [2007, February 22]. [36990]
61. Franklin, Janet; Coulter, Charlotte L.; Rey, Sergio J. 2004. Change over 70 years in a southern California chaparral community related to fire history. Journal of Vegetation Science. 15(5): 701-710. [61065]
62. Fried, Jeremy S.; Bolsinger, Charles L.; Beardsley, Debby. 2004. Chaparral in southern and central coastal California in the mid-1990s: area, ownership, condition, and change. Resource Bulletin PNW-RB-240. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 86 p. [50376]
63. Gamon, John A.; Field, Christopher B.; Goulden, Michael L.; Griffin, Kevin L.; Hartley, Anne E.; Joel, Geeske; Penuelas, Josep; Valentini, Riccardo. 1995. Relationships between NDVI, canopy structure, and photosynthesis in three Californian vegetation types. Ecological Applications. 5(1): 28-41. [63446]
64. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
65. Genin, Didier; Badan-Dangon, Antoine. 1991. Goat herbivory and plant phenology in a Mediterranean shrubland of northern Baja California. Journal of Arid Environments. 21(1): 113-121. [63324]
66. Giessow, Jason; Zedler, Paul. 1996. The effects of fire frequency and firebreaks on the abundance and species richness of exotic plant species in coastal sage scrub. In: Lovich, Jeff; Randall, John; Kelly, Mike, eds. Proceedings, California Exotic Pest Plant Council symposium; 1996 October 4-6; San Diego, CA. Volume 2. Berkeley, CA: California Exotic Pest Plant Council: 86-94. [44108]
67. Goldner, Bernard H. 1984. Riparian restoration efforts associated with structurally modified flood control channels. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of the conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 445-451. [5852]
68. Gray, John T. 1982. Community structure and productivity in Ceanothus chaparral and coastal sage scrub of southern California. Ecological Monographs. 52(4): 415-435. [3045]
69. Gray, John T. 1982. Comparative nutrient relations in adjacent stands of chaparral and coastal sage scrub. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 306-312. [6032]
70. Gray, John T. 1983. Competition for light and a dynamic boundary between chaparral and coastal sage scrub. Madrono. 30(1): 43-49. [3763]
71. Gray, John T. 1983. Nutrient use by evergreen and deciduous shrubs in southern California: I. Community nutrient cycling and nutrient-use efficiency. The Journal of Ecology. 71(1): 21-41. [63325]
72. Gray, John Timothy; Schlesinger, William H. 1981. Biomass, production, and litterfall in the coastal sage scrub of southern California. American Journal of Botany. 68(1): 24-33. [19799]
73. Green, Lisle R. 1977. Fuelbreaks and other fuel modifications for wildland fire control. Agric. Handb. 499. Washington, DC: U.S. Department of Agriculture, Forest Service. 79 p. [10511]
74. Green, Lisle R. 1981. Burning by prescription in chaparral. Gen. Tech. Rep. PSW-51. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 36 p. [19800]
75. Green, Lisle R. 1982. Prescribed burning in the California Mediterranean ecosystem. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 464-471. [6052]
76. Green, Lisle R.; Newell, Leonard A. 1982. Using goats to control brush regrowth on fuelbreaks. Gen. Tech. Rep. PSW-59. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 13 p. [10681]
77. Greenlee, Jason M.; Langenheim, Jean H. 1990. Historic fire regimes and their relation to vegetation patterns in the Monterey Bay area of California. The American Midland Naturalist. 124(2): 239-253. [15144]
78. Guo, Qinfeng. 2001. Early post-fire succession in California chaparral: changes in diversity, density, cover and biomass. Ecological Research. 16: 471-485. [42110]
79. Hall, Harvey M.; Clements, Frederic E. 1923. The phylogenetic method in taxonomy: the North American species of Artemisia, Chrysothamnus, and Atriplex. Publication No. 326. Washington, DC: The Carnegie Institute of Washington. 355 p. [43183]
80. Halligan, J. Pat. 1973. Bare areas associated with shrub stands in grassland: the case of Artemisia californica. BioScience. 23(7): 429-432. [6152]
81. Halligan, J. Pat. 1974. Relationship between animal activity and bare areas associated with California sagebrush in annual grassland. Journal of Range Management. 27(5): 358-362. [10851]
82. Halligan, J. Pat. 1975. Toxic terpenes from Artemisia Californica. Ecology. 56(4): 999-1003. [63364]
83. Halligan, J. Pat. 1976. Toxicity of Artemisia californica to four associated herb species. The American Midland Naturalist. 95(2): 406-421. [1065]
84. Halsey, Richard W. 2005. Chaparral, California's unknown wilderness. In: Halsey, Richard W. Fire, chaparral, and survival in southern California. San Diego, CA: Sunbelt Publications, Inc: 1-30. [61469]
85. Hanes, Ted L. 1971. Succession after fire in the chaparral of southern California. Ecological Monographs. 41(1): 27-52. [11405]
86. Hanes, Ted L. 1976. Vegetation types of the San Gabriel Mountains. In: Latting, June, ed. Symposium proceedings: plant communities of southern California; 1974 May 4; Fullerton, CA. Special Publication No. 2. Berkeley, CA: California Native Plant Society: 65-76. [4227]
87. Hanes, Ted L. 1977. California chaparral. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 417-469. [7216]
88. Hanes, Ted L. 1981. California chaparral. In: Di Castri, F.; Goodall, D. W.; Specht, R. L., eds. Mediterranean-type shrublands. Amsterdam: Elsevier Science Publishers B.V.: 139-174. [13576]
89. Hanes, Ted L. 1984. Vegetation of the Santa Ana River and some flood control implications. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management. Berkeley, CA: University of California Press: 882-888. [5879]
90. Hanes, Ted L.; Friesen, Richard D.; Keane, Kathy. 1989. Alluvial scrub vegetation in coastal southern California. In: Protection, management, and restoration for the 1990's: Proceedings of the California Riparian Systems conference; 1988 September 22-24; Davis, CA. Gen. Tech. Rep. PSW-110. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 187-193. [13882]
91. Harlacher, Richard A. 1985. Production of native plant materials for wildlife management programs. In: Rieger, John P.; Steele, Bobbie A., eds. Proceedings of the native plant revegetation symposium; 1984 November 15; San Diego, CA. San Diego, CA: California Native Plant Society: 62-69. [3345]
92. Harney, S. K.; Edwards, F. S.; Allen, M. F. 1997. Identification of arbuscular mycorrhizal fungi from Artemisia californica using the polymerase chain reaction. Mycologia. 89(4): 547-550. [63327]
93. Harrison, A. T.; Small, E.; Mooney, H. A. 1971. Drought relationships and distribution of two Mediterranean-climate California plant communities. Ecology. 52(5): 869-875. [6735]
94. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
95. Hield, Henry; Hemstreet, Stuart; Youngner, Victor B. 1984. Growth regulators for wild fire fuel reduction. In: Proceedings of the Plant Growth Regulator Society of America: 11th annual meeting; 1984 July 29-August 1; Boston, MA. Lagrange, GA: Plant Growth Regulator Society of America: 263-268. [63328]
96. Hillyard, Deborah; Black, Martha. 1985. Coastal sage scrub revegetation at Crystal Cove State Park. In: Rieger, John P.; Steele, Bobbie A., eds. Proceedings of the native plant revegetation symposium; 1984 November 15; San Diego, CA. San Diego, CA: California Native Plant Society: 48-52. [3344]
97. Hillyard, Deborah; Black, Martha. 1988. Coastal sage scrub revegetation at Crystal Cove State Park, Orange County, California: 1987 update. In: Rieger, John P.; Williams, Bradford K., eds. Proceedings of the second native plant revegetation symposium; 1987 April 15-18; San Diego, CA. Madison, WI: University of Wisconsin Arboretum, Society for Ecological Restoration & Management: 143-152. [4107]
98. Hobbs, E. R. 1986. Characterizing the boundary between California annual grassland and coastal sage scrub with differential profiles. Vegetatio. 65: 115-126. [1171]
99. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. [12756]
100. Horton, J. S. 1951. Vegetation. In: Some aspects of watershed management in southern California vegetation. Misc. Paper 1. Berkeley, CA: U.S. Department of Agriculture, Forest Service, California [Pacific Southwest] Forest and Range Experiment Station: 10-17. [10685]
101. Horton, J. S.; Kraebel, C. J. 1955. Development of vegetation after fire in the chamise chaparral of southern California. Ecology. 36(2): 244-262. [55799]
102. Horton, Jerome S. 1949. Trees and shrubs for erosion control of southern California mountains. Berkeley, CA: U.S. Department of Agriculture, Forest Service, California Forest and Range Experiment Station; California Department of Natural Resources, Division of Forestry. 72 p. [10689]
103. Horton, Jerome S. 1960. Vegetation types of the San Bernardino Mountains. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 29 p. [10687]
104. Horton, Jerome S.; Wright, John T. 1944. The wood rat as an ecological factor in southern California watersheds. Ecology. 25(3): 341-351. [10682]
105. Jackson, Nelroy E. 1986. Control of brush and chaparral species with glyphosate. In: Proceedings, 38th annual California weed conference; [Date Unknown]: El Macero, CA. Fremont, CA: California Weed Science Society: 221-223. [63329]
106. Kaplow, David. 1985. Coastal ecology restoration on landfill. In: Rieger, John P.; Steele, Bobbie A., eds. Proceedings of the native plant revegetation symposium; 1984 November 15; San Diego, CA. San Diego, CA: California Native Plant Society: 1-4. [3339]
107. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. In: North Carolina Botanical Garden (Producer). In cooperation with: The Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service. [36715]
108. Keeley, Jon E. 1987. Role of fire in seed germination of woody taxa in California chaparral. Ecology. 68(2): 434-443. [5403]
109. Keeley, Jon E. 1991. Seed germination and life history syndromes in the California chaparral. The Botanical Review. 57(2): 81-116. [36973]
110. Keeley, Jon E. 1995. Seed-germination patterns in fire-prone Mediterranean-climate regions. In: Arroyo, M. T. K.; Zedler, P. H.; Fox, M. D., eds. Ecology and biogeography of Mediterranean ecosystems in Chile, California, and Australia. New York: Springer-Verlag: 239-273. [48464]
111. Keeley, Jon E. 1998. Postfire ecosystem recovery and management: the October 1993 large fire episode in California. In: Moreno, J. M., ed. Large forest fires. Leiden, The Netherlands: Backbuys Publishers: 69-90. [28855]
112. Keeley, Jon E.; Fotheringham, C. J.; Baer-Keeley, Melanie. 2005. Determinants of postfire recovery and succession in Mediterranean-climate shrublands of California. Ecological Applications. 15(5): 1515-1534. [56104]
113. Keeley, Jon E.; Fotheringham, C. J.; Baer-Keeley, Melanie. 2006. Demographic patterns of postfire regeneration in Mediterranean-climate shrublands of California. Ecological Monographs. 76(2): 235-255. [63206]
114. Keeley, Jon E.; Keeley, Sterling C. 1984. Postfire recovery of California coastal sage scrub. The American Midland Naturalist. 111(1): 105-117. [5587]
115. Keller, Terry. 1993. Riparian zone plant ecology and hydrology in Aliso Creek, Chino Hills State Park, southern California. In: Keeley, Jon E., ed. Interface between ecology and land development in California: Proceedings of the symposium; 1992 May 1-2; Los Angeles, CA. Los Angeles, CA: The Southern California Academy of Sciences: 137-141. [21702]
116. Kelsey, Rick G.; Everett, Richard L. 1995. Allelopathy. In: Wildland plants: physiological ecology and developmental morphology. Denver, CO: Society for Range Management: 479-549. [27083]
117. Kirkpatrick, J. B.; Hutchinson, C. F. 1977. The community composition of Californian coastal sage scrub. Vegetatio. 35(1): 21-33. [5612]
118. Kirkpatrick, J. B.; Hutchinson, C. F. 1980. The environmental relationships of Californian coastal sage scrub and some of its component communities and species. Journal of Biogeography. 7: 23-38. [5608]
119. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384]
120. Landis, Frank C. 2000. Unburned and grazed chaparral: a case study. In: Keeley, Jon E.; Baer-Keeley, Melanie; Fotheringham, C. J., eds. 2nd interface between ecology and land development in California. U.S. Geological Survey: Open-File Report 00-62. Sacramento, CA: U.S. Department of the Interior, Geological Survey, Western Ecological Research Center: 57-69. [63307]
121. Langenheim, Jean H. 1994. Higher plant terpenoids: a phytocentric overview of their ecological roles. Journal of Chemical Ecology. 20(6): 1223-1280. [24230]
122. Little, R. John. 1981. Adventitious rooting in coastal sage scrub dominants. Madrono. 28(2): 96-97. [11030]
123. Longcore, Travis. 2003. Terrestrial arthropods as indicators of ecological restoration success in coastal sage scrub (California, U.S.A.). Restoration Ecology. 11(4): 397-409. [60029]
124. Malanson, George P. 1982. Modeling postfire succession in coastal sage scrub. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 616. [6087]
125. Malanson, George P. 1984. Fire history and patterns of Venturan subassociations of Californian sage scrub. Vegetatio. 57: 121-128. [4628]
126. Malanson, George P. 1984. Linked leslie matrices for the simulation of succession. Ecological Modelling. 21: 13-20. [6419]
127. Malanson, George P. 1985. Fire management in coastal sage-scrub, southern California, USA. Environmental Conservation. 12(2): 141-146. [1514]
128. Malanson, George P.; O'Leary, John F. 1982. Post-fire regeneration strategies of Californian coastal sage shrubs. Oecologia. 53: 355-358. [3490]
129. Malanson, George P.; O'Leary, John F. 1985. Effects of fire and habitat on post-fire regeneration in Mediterranean-type ecosystems: Ceanothus spinosus chaparral and Californian coastal sage scrub. Acta Oecologica. 6(20): 169-181. [6180]
130. Malanson, George P.; Westman, Walter E. 1985. Postfire succession in Californian coastal sage scrub: the role of continual basal sprouting. The American Midland Naturalist. 113(2): 309-318. [1516]
131. Malanson, George P.; Westman, Walter E.; Yan, Yeuh-Lih. 1992. Realized versus fundamental niche functions in a model of chaparral response to climatic change. Ecological Modelling. 64: 261-277. [48933]
132. Mayer, Audrey L.; Wirtz, William O., II. 1995. Effects of fire on the ecology of the California gnatcatcher (Polioptila californica), and associated bird species, in the coastal sage scrub community of southern California. In: Keeley, Jon F.; Scott, Tom, eds. Brushfires in California: ecology and resource management: Proceedings; 1994 May 6-7; Irvine, CA. Fairfield, WA: International Association of Wildland Fire: 77-79. [43322]
133. McBride, Joe R.; Jacobs, Diana F. 1980. Land use and fire history in the mountains of southern California. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 85-88. [16046]
134. McDonald, Philip M. 1990. Pseudotsuga macrocarpa (Vasey) Mayr bigcone Douglas-fir. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 520-526. [13412]
135. McDonald, Philip M.; Laacke, Robert J. 1990. Pinus radiata D. Don Monterey pine. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 433-441. [13401]
136. Meserve, Peter L. 1974. Ecological relationships of two sympatric woodrats in a California coastal sage scrub community. Journal of Mammalogy. 55(2): 442-447. [63415]
137. Meserve, Peter L. 1976. Food relationships of a rodent fauna in a California coastal sage scrub community. Journal of Mammalogy. 57(2): 300-319. [63395]
138. Miller, Erwin H., Jr. 1947. Growth and environmental conditions in southern California chaparral. The American Midland Naturalist. 37(2): 379-420. [63388]
139. Miller, Philip C. 1982. Nutrients and water relations in Mediterranean-type ecosystems. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 325-332. [6034]
140. Minnich, Richard A. 1976. Vegetation of the San Bernardino Mountains. In: Latting, June, ed. Symposium proceedings: plant communities of southern California; 1974 May 4; Fullerton, CA. Special Publication No. 2. Berkeley, CA: California Native Plant Society: 99-124. [4232]
141. Minnich, Richard A. 1980. Vegetation of Santa Cruz and Santa Catalina Islands. In: Power, Dennis M., ed. The California Islands: proceedings of a multidisciplinary symposium; [Date of conference unknown]; [Location of conference unknown]. Santa Barbara, CA: Santa Barbara Museum of Natural History: 123-137. [13677]
142. Minnich, Richard A. 1982. Grazing, fire, and the management of vegetation on Santa Catalina Island, California. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 444-449. [6051]
143. Minnich, Richard A. 1983. Fire mosaics in southern California and northern Baja California. Science. 219: 1287-1294. [4631]
144. Minnich, Richard A. 1999. Vegetation, fire regimes, and forest dynamics. In: Miller, P. R.; McBride, J. R., eds. Oxidant air pollution impacts in the montane forests of southern California: a case study of the San Bernardino Mountains. Ecological Studies: Analysis and Synthesis, Vol. 134. New York: Springer-Verlag: 44-80. [30370]
145. Misenhelter, Michael D.; Rotenberry, John T. 2000. Choices and consequences of habitat occupancy and nest site selection in sage sparrows. Ecology. 81(10): 2892-2901. [63378]
146. Montalvo, Arlee M.; Koehler, Catherine E. 2004. Artemisia californica. In: Francis, John K., ed. Wildland shrubs of the United States and its territories: thamnic descriptions: volume 1. Gen. Tech. Rep. IITF-GTR-26. San Juan, PR: U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry; Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 52-56. [52098]
147. Montalvo, Arlee M.; McMillan, Paul A.; Allen, Edith B. 2002. The relative importance of seeding method, soil ripping, and soil variables on seeding success. Restoration Ecology. 10(1): 52-67. [48334]
148. Montgomery, Kenneth Reid. 1976. Ether extractives and flammability of Mediterranean-type shrubs. Pomona, CA: California Polytechnic University, Pomona. 38 p. Thesis. [54096]
149. Mooney, H. A.; Dunn, E. L.; Shropshire, Frances; Song, Leo. 1970. Vegetation comparisons between the Mediterranean climatic areas of California and Chile. Flora. 159: 480-496. [13591]
150. Mooney, Harold A. 1977. Southern coastal scrub. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 471-490. [4234]
151. Moriarty, David J.; Farris, Richard E.; Stanton, Patricia A. 1985. Effects of fire on a coastal sage scrub bird community. The Southwestern Naturalist. 30(3): 452-453. [6150]
152. Moyes, Andrew B.; Witter, Martha S.; Gamon, John A. 2005. Restoration of native perennials in a California annual grassland after prescribed spring burning and solarization. Restoration Ecology. 13(4): 659-666. [60418]
153. Muller, Cornelius H. 1966. The role of chemical inhibition (allelopathy) in vegetational composition. Bulletin of the Torrey Botanical Club. 93(5): 332-351. [13672]
154. Muller, Cornelius H.; Hanawalt, Ronald B.; McPherson, James K. 1968. Allelopathic control of herb growth in the fire cycle of California chaparral. Bulletin of the Torrey Botanical Club. 95(3): 225-231. [4973]
155. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]
156. Munz, Philip A.; Keck, David D. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. [6155]
157. Myers, Marlyce A.; Ellestrand, Norman C. 1986. Post-fire succession at an inland (Riversidian) site of coastal sage scrub: variation in community response. In: DeVries, Johannes J., ed. Proceedings of the chaparral ecosystems research conference; 1985 May 16-17; Santa Barbara, CA. Report No. 2. Davis, CA: University of California, California Water Resources Center: 129-132. [4833]
158. Narog, Marcia G.; Beyers, Jan L.; Paysen, Timothy E.; Corcoran, Bonni M. 2000. Recovery of coastal sage shrub species after mechanical disturbance. In: Keeley, Jon E.; Baer-Keeley, Melanie; Fotheringham, C. J., eds. 2nd interface between ecology and land development in California. U.S. Geological Survey: Open-File Report 00-62. Sacramento, CA: U.S. Department of the Interior, Geological Survey, Western Ecological Research Center: 263-269. [63375]
159. Nord, Eamor C.; Countryman, Clive M. 1972. Fire relations. In: McKell, Cyrus M.; Blaisdell, James P.; Goodin, Joe R., eds. Wildland shrubs--their biology and utilization: Proceedings of the symposium; 1971 July; Logan, UT. Gen. Tech. Rep. INT-1. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 88-97. [1773]
160. O'Leary, John F. 1990. Californian coastal sage scrub: general characteristics and considerations for biological conservation. In: Schoenherr, Allan A., ed. Endangered plant communities of southern California: Proceedings, 15th annual symposium; 1989 October 28; Fullerton, CA. Special Publication No. 3. Claremont, CA: Southern California Botanists: 24-41. [21318]
161. Odion, Dennis C. 2000. Seed banks of long-unburned stands of maritime chaparral: composition, germination behavior, and survival with fire. Madrono. 47(3): 195-203. [38720]
162. Odion, Dennis; Tyler, Claudia. 2002. Are long fire-free periods needed to maintain the endangered, fire-recruiting shrub Arctostaphylos morroensis (Ericaceae)? Conservation Ecology. 6(2): 4. [Online]. Available: http://www.consecol.org/vol6/iss2/art4 [2006, December 15]. [63357]
163. Padgett, Pamela E.; Allen, Edith B. 1999. Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to mediterranean coastal sage scrub of California. Plant Ecology. 144(1): 93-101. [63333]
164. Parker, Robert, compiler. 1982. Reaction of various plants to 2,4-D, MCPA, 2,4,5-T, silvex and 2,4-DB. EM 4419 [Revised]. Pullman, WA: Washington State University, College of Agriculture, Cooperative Extension. 61 p. In cooperation with: U.S. Department of Agriculture. [1817]
165. Parra-Szijj, Emilia A. 1990. Revegetation in the Sepulveda Wildlife Reserve: a seven year summary. In: Hughes, H. Glenn; Bonnicksen, Thomas M., eds. Restoration '89: the new management challenge: Proceedings, 1st annual meeting of the Society for Ecological Restoration; 1989 January 16-20; Oakland, CA. Madison, WI: The University of Wisconsin Arboretum; Society for Ecological Restoration: 139-151. [14693]
166. Pase, Charles P.; Brown, David E. 1982. California coastalscrub. In: Brown, David E., ed. Biotic communities of the American Southwest--United States and Mexico. Desert Plants. 4(1-4): 86-90. [1825]
167. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
168. Peinado, M.; Alcaraz, F.; Aguirre, J. L.; Delgadillo, J. 1995. Major plant communities of warm North American deserts. Journal of Vegetation Science. 6(1): 79-94. [56028]
169. Philbrick, Ralph N., Haller, J. R. 1977. The southern California islands. In: Barbour, Michael G.; Malor, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 893-906. [7210]
170. Poole, Dennis K.; Miller, Philip C. 1975. Water relations of selected species of chaparral and coastal sage communities. Ecology. 56: 1118-1128. [10324]
171. Preston, Kris P. 1988. Effects of sulphur dioxide pollution on a Californian coastal sage scrub community. Environmental Pollution. 51(3): 179-195. [63335]
172. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
173. Rice, Elroy L. 1974. Allelopathy. New York: Academic Press, Inc. 353 p. [3317]
174. Rubinoff, Daniel. 2001. Evaluating the California gnatcatcher as an umbrella species for conservation of southern California coastal sage scrub. Conservation Biology. 15(5): 1374-1383. [63458]
175. Rundel, Philip W. 1986. Structure and function in California chaparral. Fremontia. 14(3): 3-10. [18650]
176. Sampson, Arthur W.; Jespersen, Beryl S. 1963. California range brushlands and browse plants. Berkeley, CA: University of California, Division of Agricultural Sciences; California Agricultural Experiment Station, Extension Service. 162 p. [3240]
177. Sauer, Jonathan D. 1977. Fire history, environmental patterns, and species patterns in Santa Monica mountain chaparral. In: Mooney, Harold A.; Conrad, C. Eugene, technical coordinators. Proceedings of the symposium of the environmental consequences of fire and fuel management in Mediterranean ecosystems; 1977 August 1-5; Palo Alto, CA. Gen. Tech. Rep. WO-3. Washington, DC: U.S. Department of Agriculture, Forest Service: 383-386. [4866]
178. Schultz, Brad W. 1987. Ecology of curlleaf mountain mahogany (Cercocarpus ledifolius) in western and central Nevada: population structure and dynamics. Reno, NV: University of Nevada. 111 p. Thesis. [7064]
179. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
180. Shreve, Forrest. 1927. The vegetation of a coastal mountain range. Ecology. 8(1): 27-44. [63417]
181. Smith, Robin Lee. 1980. Alluvial scrub vegetation of the San Gabriel River floodplain, California. Madrono. 27(3): 126-138. [13585]
182. Stanton, Patricia A. 1986. Comparison of avian community dynamics of burned and unburned coastal sage scrub. The Condor. 88(3): 285-289. [63372]
183. Stephens, Scott L.; Piirto, Douglas D.; Caramagno, Domenico F. 2004. Fire regimes and resultant forest structure in the native Ano Nuevo Monterey pine (Pinus radiata) forest, California. The American Midland Naturalist. 152(1): 25-36. [51279]
184. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [20090]
185. Stylinski, Cathlyn D.; Allen, Edith B. 1999. Lack of native species recovery following severe exotic disturbance in southern California shrublands. The Journal of Applied Ecology. 36(4): 544-554. [48335]
186. Taylor, Robert S., Jr. 2005. A new look at coastal sage scrub: what 70-year-old VTM plot data tell us about southern California shrublands. In: Kus, Barbara E.; Beyers, Jan L., tech. coords. Planning for biodiversity: Bringing research and management together: Proceedings of a symposium for the south coast ecoregion; 2000 February 29 - March 2; Pomona, CA. Gen. Tech. Rep. PSW-GTR-195. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 57-77. [64504]
187. Thomas, Timothy W. 1987. Population structure of the valley oak in the Santa Monica Mountains National Recreation Area. In: Plumb, Timothy R.; Pillsbury, Norman H., technical coordinators. Proceedings of the symposium on multiple-use management of California's hardwood resources; 1986 November 12-14; San Luis Obispo, CA. Gen. Tech. Rep. PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 335-340. [5384]
188. Thorne, Robert F. 1976. The vascular plant communities of California. In: Latting, June, ed. Symposium proceedings: plant communities of southern California; 1974 May 4; Fullerton, CA. Special Publication No. 2. Berkeley, CA: California Native Plant Society: 1-31. [3289]
189. Timbrook, Jan. 1990. Ethnobotany of Chumash Indians, California, based on collections by John P. Harrington. Economic Botany. 44(2): 236-253. [13777]
190. Tucker, Vance A. 1966. Diurnal torpor and its relation to food consumption and weight changes in the California pocket mouse Perognathus californicus. Ecology. 47(2): 245-252. [63389]
191. U.S. Department of Agriculture, Natural Resources Conservation Service. 2007. PLANTS Database, [Online]. Available: http://plants.usda.gov/ [2007, February 22]. [34262]
192. U.S. Department of the Interior, Fish and Wildlife Service, Division of Endangered Species. 2007. Threatened and endangered animals and plants, [Online]. Available: http://www.fws.gov/endangered/wildlife.html [2007, February 22]. [62042]
193. Vaughan, Terry A. 1954. Mammals of the San Gabriel Mountains of California. University of Kansas Publications, Museum of Natural History. Lawrence, KS: University of Kansas. 7(9): 513-582. [60582]
194. Vogl, Richard J. 1976. An introduction to the plant communities of the Santa Ana and San Jacinto Mountains. In: Latting, June, ed. Symposium proceedings: plant communities of southern California; 1974 May 4; Fullerton, CA. Special Publication No. 2. Berkeley, CA: California Native Plant Society: 77-98. [4230]
195. Vogl, Richard J. 1982. Chaparral succession. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 81-85. [6011]
196. Vogl, Richard J.; Armstrong, Wayne P.; White, Keith L.; Cole, Kenneth L. 1977. The closed-cone pines and cypress. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 295-358. [7219]
197. Walsh, Raymond C.; Reid, Thomas S. 1988. Habitat reclamation for endangered species on San Bruno Mountain. In: Rieger, John P.; Williams, Bradford K., eds. Proceedings of the second native plant revegetation symposium; 1987 April 15-18; San Diego, CA. Madison, WI: University of Wisconsin Arboretum, Society for Ecological Restoration & Management: 70-75. [4098]
198. Went, F. W.; Juhren, G.; Juhren, M. C. 1952. Fire and biotic factors affecting germination. Ecology. 33(3): 351-364. [4919]
199. Westman, W. E. 1983. Xeric Mediterranean-type shrubland associations of Alta and Baja California and the community/continuum debate. Vegetatio. 52: 3-19. [12000]
200. Westman, W. E.; O'Leary, J. F.; Malanson, G. P. 1981. The effects of fire intensity, aspect and substrate on post-fire growth of Californian coastal sage scrub. In: Margaris, N. S.; Mooney, H. A., eds. Components of productivity of Mediterranean climate regions--basic and applied aspects. The Hague, The Netherlands: Dr. W. Junk Publishers: 151-179. [13593]
201. Westman, Walter E. 1981. Diversity relations and succession in Californian coastal sage scrub. Ecology. 62(1): 170-184. [6128]
202. Westman, Walter E. 1981. Factors influencing the distribution of species of Californian coastal sage scrub. Ecology. 62(2): 439-455. [11032]
203. Westman, Walter E. 1981. Seasonal dimorphism of foliage in Californian coastal sage scrub. Oecologia. 51: 385-388. [11999]
204. Westman, Walter E. 1982. Coastal sage scrub succession. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 91-99. [6013]
205. Westman, Walter E. 1983. Island biogeography: studies on the xeric shrublands of the inner Channel Islands, California. Journal of Biogeography. 10(2): 97-118. [63427]
206. Westman, Walter E. 1985. Air pollution injury to coastal sage scrub in the Santa Monica Mountains, southern California. Water, Air, and Soil Pollution. 26(1): 19-41. [63337]
207. Westman, Walter E. 1991. Measuring realized niche spaces: climatic response of chaparral and coastal sage scrub. Ecology. 72(5): 1678-1684. [16993]
208. White, Thomas C.; Stephenson, John; Sproul, Fred. 1995. Postburn monitoring of the Eagle Fire: first year recovery on sites seeded with buckwheat and coastal sage. In: Keeley, Jon F.; Scott, Tom, eds. Brushfires in California: ecology and resource management: Proceedings; 1994 May 6-7; Irvine, CA. Fairfield, WA: International Association of Wildland Fire: 185-187. [43340]
209. Whittaker, R. H. 1970. The biochemical ecology of higher plants. In: Sondheimer, Ernest; Simeone, John B., eds. Chemical ecology. New York: Academic Press: 43-70. [12769]
210. Wiggins, Ira L. 1980. Flora of Baja California. Stanford, CA: Stanford University Press. 1025 p. [21993]
211. Wirtz, William O., II. 1974. Chaparral wildlife and fire ecology. In: Rosenthal, Murray, ed. Symposium on living with the chaparral: Proceedings; 1973 March 30-31; Riverside, CA. San Francisco, CA: The Sierra Club: 7-18. [4670]
212. Wirtz, William O., II; Mayer, Audrey L.; Raney, Mary M.; Beyers, Jan L. 1997. Effects of fire on the ecology of the California gnatcatcher, Polioptila californica, in California sage scrub communities. In: Greenlee, Jason M., ed. Proceedings, 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 91-96. [28127]
213. Yun, Kyeong W.; Maun, M. A. 1997. Allelopathic potential of Artemisia campestris ssp. caudata on Lake Huron sand dunes. Canadian Journal of Botany. 75: 1903-1912. [28273]
214. Zedler, Paul H. 1977. Life history attributes of plants and the fire cycle: a case study in chaparral dominated by Cupressus forbesii. In: Mooney, Harold A.; Conrad, C. Eugene, technical coordinators. Symposium on the environmental consequences of fire and fuel management on Mediterranean ecosystems: Proceedings; 1977 August 1-5; Palo Alto, CA. Gen. Tech. Rep. WO-3. Washington, DC: U.S. Department of Agriculture, Forest Service: 451-458. [4876]
215. Zedler, Paul H. 1981. Vegetation change in chaparral and desert communities in San Diego County, California. In: West, D. C.; Shugart, H. H.; Botkin, D. B., eds. Forest succession: concepts and application. New York: Springer-Verlag: 406-430. [4241]
216. Zedler, Paul H. 1982. Plant demography and chaparral management in southern California. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 123-127. [6016]
217. Zedler, Paul H.; Gautier, Clayton R.; McMaster, Gregory S. 1983. Vegetation change in response to extreme events: the effect of a short interval between fires in California chaparral and coastal scrub. Ecology. 64(4): 809-818. [4612]
218. Zink, Thomas A.; Allen, Michael F. 1998. The effects of organic amendments on the restoration of a disturbed coastal sage scrub habitat. Restoration Ecology. 6(1): 52-58. [63340]

FEIS Home Page