SPECIES: Ceanothus cuneatus

• Introductory
• Distribution and occurrence
• Botanical and ecological characteristics
• Fire ecology
• Fire effects
• Management considerations
• References

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION SYNONYMS NRCS PLANT CODE COMMON NAMES TAXONOMY LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	© Michael W. Tuma

Ceanothus cuneatus var. cuneatus (buckbrush)
Ceanothus cuneatus var. fascicularis (McMinn) Hoover (sedgeleaf buckbrush) [55,58]
Ceanothus cuneatus var. rigidus (Nutt.) Hoover (Monterey ceanothus) [55,58]

Hybrids: Ceanothus distribution shares a wide range of habitats and distributions with other Ceanothus species and is thought to hybridize frequently [55,78]. Several hybrids are recognized in older floras. Current floras do not recognize any of these hybrids as separate species.

LIFE FORM:
Shrub

FEDERAL LEGAL STATUS:
None

OTHER STATUS:
None

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES/PROVINCES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• HABITAT TYPES AND PLANT COMMUNITIES

Plants database provides a distributional map of Wedgeleaf ceanothus and its infrataxa.

ECOSYSTEMS [42]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES28 Western hardwoods
FRES29 Sagebrush
FRES34 Chaparral-mountain shrub

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

CA

OR

MEXICO

B.C.N.

B.C.S.

BLM PHYSIOGRAPHIC REGIONS [14]:
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains

KUCHLER [69] PLANT ASSOCIATIONS:
K002 Cedar-hemlock-Douglas-fir forest
K005 Mixed conifer forest
K009 Pine-cypress forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K024 Juniper steppe woodland
K026 Oregon oakwoods
K029 California mixed evergreen forest
K030 California oakwoods
K033 Chaparral
K034 Montane chaparral
K035 Coastal sagebrush
K036 Mosaic of K030 and K035
K047 Fescue-oatgrass
K048 California steppe
K055 Sagebrush steppe

SAF COVER TYPES [36]:
210 Interior Douglas-fir
229 Pacific Douglas-fir
233 Oregon white oak
234 Douglas-fir-tanoak-Pacific madrone
237 Interior ponderosa pine
238 Western juniper
241 Western live oak
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
248 Knobcone pine
249 Canyon live oak
250 Blue oak-foothills pine
255 California coast live oak

SRM (RANGELAND) COVER TYPES [98]:
107 Western juniper/big sagebrush/bluebunch wheatgrass
110 Ponderosa pine-grassland
109 Ponderosa pine shrubland
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
209 Montane shrubland
214 Coastal prairie
215 Valley grassland
216 Montane meadows
405 Black sagebrush

HABITAT TYPES AND PLANT COMMUNITIES:
Wedgeleaf ceanothus is commonly encountered in most chaparral vegetation types and several forest communities throughout California, Oregon and the Baja of Mexico. While sometimes found in pure stands representing the dominant vegetation, wedgeleaf ceanothus more often codominates or associates with other species in shrub stands or is a substantial understory species of pine (Pinus spp.) forests or oak (Quercus spp.) woodlands [30].

In California wedgeleaf ceanothus is the dominant shrub species in the ceanothus chaparral vegetation type. Shrub species that may associate with wedgeleaf ceanothus include chamise (Adenostoma fasciculatum), hoaryleaf ceanothus (Ceanothus crassifolius), hairy ceanothus (C. oliganthus), blueblossom (C. thyrsiflorus), Nuttall's scrub oak (Q. dumosa), toyon (Heteromeles arbutifolia), and sugar sumac (Rhus ovata) [52].

Chamise chaparral is the most common type of chaparral in California occurring in the north and central Coast Ranges, Sierra Nevada foothills, southern California and northern Baja mountain ranges. This type of chaparral is usually dominated by chamise, although in many stands, wedgeleaf ceanothus codominates with chamise and/or whiteleaf manzanita (Arctostaphylos viscida) [7]. Stands where chamise and wedgeleaf ceanothus codominate are sometimes referred to as mixed chaparral. Species that associate with wedgeleaf ceanothus in this cover type include trees such as blue oak (Q. douglasii) and California buckeye (Aesculus californica), shrubs such as red shank (Adenostoma sparsifolium), Nuttall's scrub oak, birchleaf mountain-mahogany (Cercocarpus betuloides), laurel sumac (Malosma laurina), white and black sage (Salvia mellifera, S. apiana), sugar sumac, Our Lord's candle (Yucca whipplei), and herbs such as giant wildrye (Leymus condensatus), and Eastern Mojave buckwheat (Eriogonum fasciculatum) [8,18,52,74].

The most diverse community where wedgeleaf ceanothus frequently occurs in is the montane chaparral of the lower elevations and xeric sites of the Cascade, Klamath, and Siskiyou mountains of southwestern Oregon and northern California, the Transverse and Peninsular ranges of southern California, and the Sierra San Pedro Mártir of northern Baja. Habitat types in this category are foothill woodlands, and mixed coniferous forest. Generally this cover type refers to occurrences of wedgeleaf ceanothus found in the understory of transmontane forested slopes of Jeffrey pine (Pinus jeffreyi) and gray pine (P. sabiniana) in California, and Pacific ponderosa pine (P. ponderosa var. ponderosa) and oak woodlands of California and Oregon. Characteristic species that associate with wedgeleaf ceanothus in this cover type include trees such as Oregon white oak (Q. garryana), blue oak, California black oak (Q. kelloggii), California shrub live oak (Q. turbinella var. californica), valley oak (Q. lobata), leather oak (Q. durata), interior live oak (Q. wislizenii), coast live oak (Q. agrifolia), canyon live oak (Q. chrysolepis), and California buckeye. Shrubs include whiteleaf manzanita, bigberry manzanita (Arctostaphylos glauca), yerba santa (Eriodictyon californicum), eastern redbud (Cercis canadensis), pointleaf manzanita (A. pungens), Klamath plum (Prunus subcordata), California buckthorn (Frangula californica ssp. cuspidata), common snowberry (Symphoricarpos albus), Mojave ceanothus (Ceanothus greggii var. vestitus), Mohave buckbrush (C. g. var. perplexans), birchleaf mountain-mahogany, thickleaf yerba santa (E. crassifolium), flannelbush (Fremontodendron californicum), California coffeberry (Rhamnus californica), yellowleaf silktassel (Garrya flavescens), and poison-oak (Toxicodendron diversilobum) [19,30,35,52,57,71,84,96]. Also in the montane chaparral, wedgeleaf ceanothus associates with less frequented stands of Baker cypress (Cupressus bakeri) in northern California and southern Oregon [31,101], Tecate cypress (C. forbesii) in southern California and Baja [5,32], and bigcone Douglas-fir (Pseudotsuga macrocarpa) in southern California mountains [43]. In Siskiyou County, California, and on lava flows in eastern Shasta County, California, wedgeleaf ceanothus associates with small populations of western juniper (Juniperus occidentalis) [23].

The coastal sage scrub habitat type is dominated by California sagebrush (Artemisia californica) and includes wedgeleaf ceanothus in areas in or near low elevation coastal aspects. Other species that may associate with wedgeleaf ceanothus in this cover type include white, black, and purple sage (Salvia leucophylla), California brittlebush (Encelia californica), eastern Mojave buckwheat, and thickleaf yerba santa [34,52,75].

In California small populations of wedgeleaf ceanothus are found on inland dune locations which have minimal soil development. Species that commonly associate with wedgeleaf ceanothus in these communities include coast live oak, chamise, California buckeye, Santa Barbara ceanothus (Ceanothus impressus), California prickly phlox (Leptodactylon californicum), and black sage [9].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

Wedgeleaf ceanothus is a native, perennial, evergreen shrub reaching heights of 3.3 to 11.5 feet (1-3.5 m) tall. The branches are rigid. Leaves are opposite, firm, flat, and 5 to 15 mm long, although considerable differences in leaf size have been observed, which is thought to be driven by water availability [27]. There is evidence that wedgeleaf ceanothus can withdraw nutrients from senescing leaves [86]. For more information see Seedling establishment/growth. Flowers are 5 to 6 mm broad with short erect horns near the top. Fruits are capsules that contain 2 to 3 seeds, round to oblong in shape, and 0.16 inches (4 mm) long [13,35,55,56,58,78,109]. Roots are many branched from a single tap root and can penetrate "deeply" into the soil [96].

Wedgeleaf ceanothus establishment is generally synchronous after burning so wedgeleaf ceanothus stands are usually even-aged [60]. Biswell [21] feels substantial mortality of wedgeleaf ceanothus begins in stands more than 50 years old.

RAUNKIAER [88] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Wedgeleaf ceanothus regenerates from seed [59].

Breeding system: Wedgeleaf ceanothus is monoecious [58].

Pollination: Wedgeleaf ceanothus is cross-pollinated by insects [58].

Seed production: Wedgeleaf ceanothus seed production varies yearly [22].

Seed dispersal: occurs during the spring [61]. The mature capsule bursts upon opening, making an audible pop, and seeds are cast up to a distance of 35 feet (10.7 m) [22]. However, the majority of seeds fall near the parent shrub [35]. Seed casting date and distance depend on phenology of fruit-ripening, temperature, and humidity. Hotter and drier conditions result in further casting which generally occurs during the hot and dry months of July and August [35].

Seed can also be dispersed by insects. California harvester ants are responsible for caching a considerable amount of wedgeleaf ceanothus seed below ground, which is thought to protect seeds from lethal temperatures during burning [79].

Seed banking: Seeds of wedgeleaf ceanothus are hard-coated, nearly impermeable, and may lie viable in the ground for many years [22,61]. Viable wedgeleaf ceanothus seed are commonly found buried in soils of chaparral [102]. Exactly how long banked wedgeleaf ceanothus seed can remain viable needs to be investigated.

Germination: occurs during the spring following fire [102]. Wedgeleaf ceanothus seeds require relatively high temperatures during burning (158 to 212 °F (70-100 °C) to facilitate germination [90]. Germination rates are high after fire [33,96] which scarifies wedgeleaf ceanothus seed [87,102]. Heat from fire melts or cracks the cuticle of buried seeds [61] which is necessary for germination. Sweeney [102] investigated the effects of fire on seed, and found a majority of wedgeleaf ceanothus seed germinated after being exposed to varying degrees of temperature up to 176 °F (80 °C). The effects of higher temperatures are unknown [102]. Germination rates are significantly (P<.01) enhanced when charate (chemical release from burnt wood) from chamise was used synergistically with heat in germination experiments [59]. The mechanism behind charate-stimulated germination is unknown. While germination is stimulated by burning, in the absence of fire, wedgeleaf ceanothus can germinate in shrub overstory openings [18,22]. In a greenhouse environment germination of wedgeleaf ceanothus was most successful when seeds were planted at depths of 0.5 to 1 inch (1.3-2.5 cm) [2,13].

Seedling establishment/growth: Wedgeleaf ceanothus is widely considered an "obligate seeder" or "fire-recruiter." Regeneration depends almost entirely on germination from seed during postfire conditions [1,20,62,102]. During the spring after burning, varying numbers of wedgeleaf ceanothus seedlings appear. Very high mortality rates are common during the 1st year after establishment. This is believed to be caused by summer drought and interference from herbaceous competitors (see Plant Response To Fire) [33,96,102]. Schultz and others [96] reported the emergence of wedgeleaf ceanothus seedlings in the central California Sierra Nevada foothills occurred in mid-March and April. By mid-June, root depths may reach 30 to 40 inches (76-102 cm) while above ground stems and branches may reach 6 to 8 inches (15-20 cm) tall. Wedgeleaf ceanothus roots penetrate much further than those of herbaceous competitors. It is believed that wedgeleaf ceanothus' vigorous root growth beyond the maximum penetration of grass roots in the 1st year of growth is critical to obtaining enough moisture to establish [96]. Wedgeleaf ceanothus in the absence of fire may become established in shrub openings [22]. In their review of chaparral vegetation, Keeley and Keeley [65] point out that obligate-seeding shrubs, including wedgeleaf ceanothus, have more opportunities for genetic recombination than obligate-sprouting species. Recruitment of wedgeleaf ceanothus between fire intervals is very uncommon and rarely results in successful establishment under the canopy of mature shrubs, even in stands unburned for more than a century [60,61].

Asexual regeneration: The ability of wedgeleaf ceanothus to regenerate vegetatively is unclear. One observation from Biswell and Gilman [22] noted asexual regeneration of wedgeleaf ceanothus through layering after a prolonged period without fire and grazing.

SITE CHARACTERISTICS:
Wedgeleaf ceanothus covers a wide array of geographic and topographic locations from valley floors to hillsides and foothill slopes. It generally occurs in elevations < 6000 feet (1800 m) in California and Oregon on dry mountain slopes and ridges within the Upper Sonoran Life Zone [55].

Climate of this region is considered "Mediterranean" with a majority of annual precipitation occurring in winter with long summer droughts. Typically wedgeleaf ceanothus occurs in areas where annual precipitation ranges from approximately 10 to 35 inches (250-900 mm) and where 80% of the annual total precipitation occurs in the fall, winter, and spring [30]. Annual average precipitation ranges from north to south:

State	Location	Mean Annual Precipitation	Citation
Oregon	Medford	16.5 inches (419 mm)	[30]
California	Santa Rosa	35 inches (888.5 mm)	[25]
California	Fresno	29.9 inches (760 mm)	[35]
California	San Mateo County	25.7 inches (654 mm)	[1]
California	Los Angeles	15.7 inches (400 mm)
California	San Diego	10 inches (250 mm)	[77]

Wedgeleaf ceanothus occurs in chaparral vegetation types in California and Oregon and is commonly associated with poor, rocky soils [41,61]. Wedgeleaf ceanothus is more frequently found growing on nonserpentine soils of sandstone origins than on serpentine soils [92]. However, wedgeleaf ceanothus can be found on both types of soils and is considered an indicator species for field identification of serpentine soil conditions in California and Oregon [68].

SUCCESSIONAL STATUS:
Wedgeleaf ceanothus stand development is not easily predicted and is dependent on a number of variables including plant association, habitat type, proximity to boundaries with other habitat types, geographic and topographic location, climate, fire intensity, and time since last fire [51]. This is in contrast to early theories that outline succession in chaparral as an orderly progression of seral stages that reach a climax as originally defined by Clements [91].

Wedgeleaf ceanothus stands change rapidly during the first 1 to 4 years postfire. In areas where wedgeleaf ceanothus associates with sprouting shrub species postfire succession can typically be described in 3 stages: (1) During the 1st postfire year native and nonnative vegetation forms the dominant cover, while chaparral shrub seedlings and sprouts emerge. (2) During the 2nd postfire year, high mortality of shrub and subshrub seedlings takes place with decreased native and increased nonnative herbaceous plants. (3) In subsequent years, the remaining shrub seedlings and sprouts become well established while herbaceous vegetation gradually decreases. After 8 to 10 years, a relatively mature chaparral cover with little understory exists [52,102]. Very little is known about the average life span of wedgeleaf ceanothus, although many agree with Biswell's [21] observations that substantial mortality begins in stands >50 years old.

Stand development seems to be largely driven by water availability. While sometimes found in pure stands, wedgeleaf ceanothus often codominates or associates with other shrub species during stand development. The subsequent dominance of any one species is greatly influenced by water availability and can be a major contributor to the occurrence and frequency of wedgeleaf ceanothus in the resulting stand structure [40].

In most stands wedgeleaf ceanothus can form impenetrable thickets that may retard understory development of other plant species [26]. However, in long disturbance-free periods, wedgeleaf ceanothus stands can undergo decline because of interference from introduced sprouting shrubs or overstory species from nearby stands [40]. For example, suppression of fire in chaparral is thought to favor crown-sprouting species over obligate seeders. Research conducted in the southern coastal ranges of California found that fire suppression is primarily responsible for the conversion of large acreages of shrub lands where wedgeleaf ceanothus occurred to oak woodlands [106]. Wedgeleaf ceanothus has shown significant decreases in areas susceptible to shading from overstory species, in particular by Nuttall's scrub oak and toyon on coastal ranges [63,106]. Generally wedgeleaf ceanothus declines in undisturbed stands that reach > 100 years old [53,60]. In unburned areas of the south coastal ranges of California [20], stands of wedgeleaf ceanothus that have not burned for over a century are replaced by longer lived species such as chamise [40].

Wedgeleaf ceanothus is an actinorrhizal plant that has the ability to fix atmospheric nitrogen [24,28,29]. This gives wedgeleaf ceanothus a competitive advantage over other non-nitrogen fixing shrubs herbs and grasses, especially on nitrogen-deficient soils [24]. Over a given year wedgeleaf ceanothus nodulates nitrogen at an estimated rate of 54 pounds per acre [29].

SEASONAL DEVELOPMENT:
Wedgeleaf ceanothus occurs in the Mediterranean-climate zone where annual summer drought is typical. The unpredictability of both intensity and duration of this drought has a major influence on the development strategies of wedgeleaf ceanothus. All plant growth must occur before water stress triggers dormancy. A 2-year study in the foothills of Sequoia National Park, California, found that wedgeleaf ceanothus begins phenological development in late winter and early spring and exhibits simultaneous branch elongation, leaf initiation, and flowering. This adaptation insures completion of all phenological stages before the onset of drought [7]. Wedgeleaf ceanothus is able to survive extreme drought conditions as observed during the 1975-1977 drought in California [81].

Wedgeleaf ceanothus flowers from February to April depending on location [109]. Leaf life span averaged 14.4 months in the eastern foothills of the Santa Cruz Mountains [1]. Wedgeleaf ceanothus thrives in the cool, wet winter and withers during the dry summer. Wedgeleaf ceanothus is considered a sclerophyll which is characterized by small leaves, short internodes, thick cuticle, sunken stomates, high proportion of lignified cells, and leaves with a waxy coating. All of these traits help wedgeleaf ceanothus to survive water loss through transpiration [1].

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Fire regimes: Opinions among chaparral scientists conflict on the degree to which chaparral is dependent on fire [40]. Historical fire intervals of 30 to 100 years appear most favorable for wedgeleaf ceanothus stand maintenance [77,82]. Theoretically, longer fire intervals favor wedgeleaf ceanothus by allowing larger quantities of annually-deposited, long-lived seed to accumulate. This provides better chances for postfire establishment [80]. This is counter to many land management fire prescriptions, especially in wildland-urban interface areas, where hazardous fuel reduction is a priority. Human caused ignitions, intentional and unintentional, cause fire intervals of 20-30 years, especially in stands in close proximity to towns or cities [60]. These intervals may be too short for sufficient seed accumulation.

Burning by Native Americans: Before European settlement, burning by Native Americans impacted fire intervals and vegetation structure, especially in areas where wedgeleaf ceanothus occurs [4]. This is especially apparent in chaparral stands or oak woodlands where wedgeleaf ceanothus and other chaparral shrub species are common in the understory. In California, observations in oak woodlands in the mid-twentieth century found increasing densities of chaparral species, including wedgeleaf ceanothus, in the understory of oaks. This is believed to be due to suppression of native American burning practices and wildfire [17]. Burning by aboriginals in California was thought to be primarily for maintenance of hunting grounds and prevention of large "devastating" fires in mature stands of wedgeleaf ceanothus, a species recognized by natives to be very important to wildlife. Native Americans probably ignited low-intensity grassfires during the spring in oak woodlands and winter range of regional ungulate species, to prevent wedgeleaf ceanothus from being consumed by intense, naturally-ignited fires that typically occurred during mid-summer [17,73]. The extent and rationale of burning in chaparral by natives is in need of further study, especially in the context of ecological restoration.

The following table provides fire return intervals for plant communities and ecosystems where wedgeleaf ceanothus is important. It may not be inclusive. If you are interested in plant communities or ecosystems that are not listed, see the complete FEIS Fire Regime Table.

Community or Ecosystem	Dominant Species	Fire Return Interval Range (years)
California chaparral	Adenostoma and/or Arctostaphylos spp.	< 35 to < 100
coastal sagebrush	Artemisia californica	< 35 to < 100
California montane chaparral	Ceanothus and/or Arctostaphylos spp.	50-100 [82]
California steppe	Festuca-Danthonia spp.	< 35 [82,100]
western juniper	Juniperus occidentalis	20-70 [82]
Jeffrey pine	Pinus jeffreyi	5-30
Pacific ponderosa pine*	Pinus ponderosa var. ponderosa	1-47
California oakwoods	Quercus spp.	< 35 [6]
coast live oak	Quercus agrifolia	2-75 [50]
coastal Douglas-fir*	Pseudotsuga menziesii var. menziesii	40-240 [6,54,91]
California mixed evergreen	Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii	<35
canyon live oak	Quercus chrysolepis	<35 to 200
blue oak-foothills pine	Quercus douglasii-P. sabiniana	<35
Oregon white oak	Quercus garryana	< 35 [6]
California black oak	Quercus kelloggii	5-30 [82]
interior live oak	Quercus wislizenii	< 35 [6]

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

POSTFIRE REGENERATION STRATEGY [99]:
Shrub without adventitious bud/root crown
Ground residual colonizer (on-site, initial community)
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Since wedgeleaf ceanothus establishment is synchronous after burning, wedgeleaf ceanothus typically forms even-aged stands [60]. Wedgeleaf ceanothus stands can become very dense at maturity [22]. Frequently during early stages of wedgeleaf ceanothus stand development, stands are densely crowded and allow few, if any, herbaceous species to occur under the canopy [22]. As stands mature, self-thinning occurs; then stand structure, density, and composition stabilize. Stand thinning seems to result from intraspecific competition for moisture and is highly correlated with how close shrubs grow to one another. Clumped distributions tend to show the greatest rates of mortality during times of high moisture stress. This pattern of mortality is present in both juvenile and mature stands of wedgeleaf ceanothus and other ceanothus species [63].

Postfire establishment patterns at small scales: Wedgeleaf ceanothus stands can become very dense during maturity allowing few, if any, herbaceous species to occur under its canopy [22]. In these areas postfire emergence and survival of seedlings is high due to a lack of interference from grasses and other herbaceous species [22]. Florence and Florence's [39] observations from prescribed fire activities noted that postfire wedgeleaf ceanothus seedlings were frequently found near burned skeletons of wedgeleaf ceanothus or other sprouting shrubs. They assumed that the dead remains of the shrubs provided a better habitat for the wedgeleaf ceanothus seedlings by protecting the seedlings from browsing. Also the authors hypothesized that high temperatures or charate released during burning may have enhanced germination of wedgeleaf ceanothus by increasing mortality of other competing herbaceous and shrub species [39].

Fire and nodulation of wedgeleaf ceanothus: In California an irrigation experiment during mid-summer investigated whether water stress inhibits nodulation of postfire wedgeleaf ceanothus seedlings. The authors found a significant (P< 0.05) increase in nodulation frequency in well-hydrated sites, compared to adjacent xeric sites, which are typical of wedgeleaf ceanothus habitat. Also noted in this study was an interesting delay in nodulation in 1st year postfire seedlings. Nodulation of postfire seedlings did not occur at the onset of spring immediately after germination when soil moisture values where high. Pratt and others [85] suggested that wedgeleaf ceanothus might be able to suppress nodulation by delaying nitrogen fixation until adequate carbohydrate reserves and water availability are synchronously established.

FIRE MANAGEMENT CONSIDERATIONS:
Wildfire suppression: policies during the 20th century have interrupted the natural fire cycle of many types of chaparral including those where wedgeleaf ceanothus occurs. Current management of chaparral stands includes the use of prescribed fire to maintain natural fire regimes. However, large areas of continuous "decadent" wedgeleaf ceanothus chaparral exist where fire has not occurred for over a century. Fires in these areas burn with high intensity over large areas, potentially beyond historical levels of severity [38]. Consequently, fire suppression's overall effects are thought to reduce the numbers of wildfires, but increase overall area burned [76,107]. Note that wedgeleaf ceanothus stems identified as "decadent" may not necessarily be dead; see [59].

Frequency of burning: Research conducted during the mid-20th century focused on using frequent fires to reduce wedgeleaf ceanothus and chamise stands for browse habitat improvements (see Importance To Livestock And Wildlife). Stands of wedgeleaf ceanothus can be decimated when fires are frequent enough to kill postfire seedlings that have not matured enough to produce a seed crop [20]. One experimental burn conducted 3 years after burning on a young stand of wedgeleaf ceanothus resulted in 100% mortality of all stems and 0% postfire establishment of new wedgeleaf ceanothus seedlings the following spring [53]. This phenomenon has been called "shock stagnation," a semipermanent degradation of the native vegetation in which exotic grasses and/or forbs dominate. This tactic was frequently used during the mid-1900s [59].

The duration of time that wedgeleaf ceanothus can exist without fire is unknown.

Seasonality of Burning: Wedgeleaf ceanothus seedlings must develop considerable root systems during the spring before the cessation of seasonal rains. Middle- to late-spring burns may result in very high wedgeleaf ceanothus seedling mortality [20,39].

Wedgeleaf ceanothus exhibited successful rates of establishment when burning was conducted before winter [39]. The authors believe burning before the cool season allows seedlings of wedgeleaf ceanothus to establish before annual herbaceous species arrive, giving wedgeleaf ceanothus a competitive moisture advantage and subsequent higher rates of survival through the following dry season [39]. However, low fire intensities commonly associated with prescribed fires during the cool season are also directly correlated with high coverages of herbaceous species during the 1st year postfire [38]. Most prescribed fires occur during the cool season due to safety issues. It may be difficult to use prescribed fire for ecological restoration while addressing safety needs. Further research is needed on the effects of seasonality of burning and establishment wedgeleaf ceanothus.

Postfire establishment and interference: One concern following fire is possible interference from herbaceous species. Postfire mortality of wedgeleaf ceanothus seedlings suggests competition of water and nutrient resources from nearby herbaceous plants is an important factor influencing initial survival rates [102]. This effect may be less common in areas with low rainfall. Schultz and others [96] in the Sierra Nevada of central California found the abundance and vigor of wedgeleaf ceanothus seedlings were negatively associated with increasing densities of herbaceous species, especially grasses. Postfire establishment of grasses in this area usually precedes wedgeleaf ceanothus, quickly creating a mass of roots difficult for shrub species to push young roots through. The authors believed competition between grasses and wedgeleaf ceanothus seedlings for moisture during establishment is responsible for dramatic reductions in numbers of wedgeleaf ceanothus seedlings 1 year postfire. In their study, 3 months after emergence, wedgeleaf ceanothus growing without interference from other species developed roots to a depth of 43 inches (109 cm) and had 26 inches (66 cm) of lateral growth. In contrast, under the same watering regime, wedgeleaf ceanothus seedlings growing along with Italian ryegrass (Lolium multiflorum) had a maximum root depth of 11.5 inches (29 cm) with very little lateral growth [96]. Wedgeleaf ceanothus seedling mortality was highest when density of Italian ryegrass was >39% at maturity. Biswell [21] found that native herbaceous density > 65% severely affected survival of wedgeleaf ceanothus and other chaparral shrubs. Contrary to Schultz and others [96], Beyers and others [15] found no significant (P>0.05) differences in wedgeleaf ceanothus stem densities in plots in southern California that either had or had not been reseeded with Italian ryegrass 5 years postfire. However in this study grass densities were low and rainfall less than average [96].

Postfire competition for light often controls shrub species dominance in chaparral. In ecotonal regions between coastal sage and chaparral, stands 30 years postfire showed declining density of purple sage in mixed stands with wedgeleaf ceanothus. This was not caused by allelopathy but by competition for light. During stand maturity wedgeleaf ceanothus grows taller and shades purple sage [75]. Similar results have been found from comparable research using different Ceanothus species. [94].

Fire intensity and postfire establishment: Wedgeleaf ceanothus establishment is associated with areas where the prefire canopy was dense and consequently high fire severity occurred. Soil heating is the primary trigger to end dormancy for wedgeleaf ceanothus (see Seedling establishment/growth). A study hypothesized that in the absence of fire, abnormally large accumulations of fuel over a period of time would result in extreme fire severities, reducing numbers of buried viable seed [76]. However, the seed of wedgeleaf ceanothus was found to be very resistant to heat and high fire severity [80]. Prescribed fires in chaparral types are conducted during the fall after the onset of the rainy season and generally exhibit lower fire intensity than typical wildfires during late summer or early fall [33]. Effects of low intensity burns on germination rates of wedgeleaf ceanothus are not well known.

Most fire ecology studies conducted in chaparral vegetation have focused attention on the fire ecology of stands that exist on sandstone-derived soils, leaving the fire ecology of stands on serpentine soils largely uninvestigated. Wedgeleaf ceanothus is found in serpentine soils, though it is more common and more abundant on nonserpentine soils [92]. This could be due to lower fire severities. Serpentine soils generally support lower densities of chaparral species, have lower concentrations of fuel, and burn with lower severity than fires on nonserpentine soils. These fires may not break seed dormancy in wedgeleaf ceanothus [92]. For more information on serpentine flora ecology see [68].

Pre-and postfire grazing: Timing and intensity of cattle grazing can affect wedgeleaf ceanothus. Heavy browsing of wedgeleaf ceanothus lowers seed production and reduces the potential for future establishment of wedgeleaf ceanothus after burning. In areas where heavy grazing occurs, young wedgeleaf ceanothus seedlings may not produce sufficient seed before being grazed to regenerate after fire [22]. The combined effects of burning and heavy grazing on wedgeleaf ceanothus have been used to convert stands of wedgeleaf ceanothus to pasture [21]. Biswell and Gilman [22] recommended burning in areas susceptible to heavy grazing at intervals >20 years to allow for sufficient stand development and seed production.

Wedgeleaf ceanothus is especially sensitive to browsing by deer after fire. Light to heavy browsing on young wedgeleaf ceanothus seedlings has reduced abundance of seedlings following burning [20]. In California, after an unknown amount of time after burning, "light" browsing by deer resulted in stunted seedlings averaging heights of 18 inches (46 cm). Seedlings protected from browsing by enclosures grew rapidly and averaged >27 inches (69 cm) in height. Mortality rates of stems from browsing were dramatically reduced after shrubs reached 5 years of age [20].

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

After fire, deer and other ungulates prefer grazing in postfire stands of wedgeleaf ceanothus. Peak browsing in these areas occurs for up to 3 years after burning [66,67].

Effects of cattle grazing in stands of wedgeleaf ceanothus along with manipulation of chaparral for "range improvement" and "improvement of wildlife habitat" are well documented. For more information please refer to [22,44,45,46,47,49,93,95,103].

Palatability/nutritional value: The foliage, twigs, and seedlings of wedgeleaf ceanothus are highly palatable to mule deer, black-tailed deer, and domestic sheep and goats [20,47,97]. Overall palatability to cattle is low [104]. Seeds are highly palatable to many small mammals, birds, and insects [26].

Wedgeleaf ceanothus offers year around high-protein browse for black-tailed deer, mule deer and other wildlife species [16]. Domestic sheep prefer wedgeleaf ceanothus for browse, while cattle will eat wedgeleaf ceanothus when other forage is scarce [22]. The following table shows monthly fluctuations in crude protein content of wedgeleaf ceanothus [16]. Values equal percentages of crude protein from oven-dried plant material. Crude protein peaks in spring and summer.

January	February	March	April	May	June	July	August	September	October	November	December
8.0	7.8	9.2	15.6	12.4	9.5	7.9	10.4	7.5	6.5	7.0	8.1

Wedgeleaf ceanothus provides high levels of nutrients important to ungulate species. Mineral concentrations of wedgeleaf ceanothus for healthy deer populations are reported by Scrivner and others [97]. The following table shows mean percent mineral composition of oven-dry wedgeleaf ceanothus material sampled from June, 1985, to July, 1986, in California.

P	S	Ca	Mg	K
0.12	0.11	0.62	0.23	0.74

Gordon and Sampson [48] also provide nutritional information on wedgeleaf ceanothus.

Cover value: Wedgeleaf ceanothus provides cover for many wildlife species including California quail, black-tailed jackrabbit, brush rabbit, and mourning dove [20,26]. The preferred habitat of the chaparral mouse is under the protective branches of wedgeleaf ceanothus [70]. Many other small rodents including the deer mouse, California mouse, house mouse, and California pocket mouse, hide, feed, and nest beneath the canopy of wedgeleaf ceanothus [70]. Plants frequently grow tall enough, and with sufficient density, to furnish good hiding cover for larger ungulates such as mule deer. Barrett [10] found that black-tailed deer preferred wedgeleaf ceanothus chaparral over other adjacent cover types for browse in the foothill region of Mt Lassen, California.

VALUE FOR REHABILITATION OF DISTURBED SITES:
Wedgeleaf ceanothus is well suited for use in rehabilitation because of rapid growth rates and an ability to improve soil fertility through nitrogen fixation. Some cultivars are now commercially available [37]. Wedgeleaf ceanothus has been successfully planted onto many types of disturbed sites throughout southern California and the desert Southwest [37]. It established well on disturbed sites near Lake Tahoe, California, but exhibited poor long-term survival due to cold winter temperatures [99]. Properly treated seed can be hand-sown onto burned slopes as an emergency revegetation measure in southern California chaparral. Good seedling establishment has been reported following seeding of these sites [12]. Wedgeleaf ceanothus can be used for stabilization of neutral and acid soils. However, transplanting from a nursery is recommended due to the difficulty and expense of harvesting seeds [83].

OTHER USES:
The Miwok Indians of the Sierra Nevada region of California used the young, straight shoots of wedgeleaf ceanothus for basketry material. The young shoots are so valuable that the Miwok have historically manipulated stands of wedgeleaf ceanothus by pruning, burning, or coppicing to induce rapid elongation of young growth [3]. The Kawaiisu used straightened twigs of wedgeleaf ceanothus for arrows and also used the shrub for fire wood [110]. The Mono tribe used stems of wedgeleaf ceanothus for basketry materials [4].

OTHER MANAGEMENT CONSIDERATIONS:
No information is available for this topic.

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