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Ceanothus cordulatus



INTRODUCTORY


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AUTHORSHIP AND CITATION:
Reeves, Sonja L. 2006. Ceanothus cordulatus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/shrub/ceacor/all.html [].

FEIS ABBREVIATION:
CEACOR

SYNONYMS:
None

NRCS PLANT CODE [100]:
CECO

COMMON NAMES:
whitethorn ceanothus
mountain whitethorn
snow bush

TAXONOMY:
The scientific name of whitethorn ceanothus is Ceanothus cordulatus Kellogg (Rhamnaceae) [51]. Intrasectional hybridization within Ceanothus species is common [67]. When information specific to whitethorn ceanothus is not available, references will be made to the genus Ceanothus.

LIFE FORM:
Shrub

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
None

DISTRIBUTION AND OCCURRENCE

SPECIES: Ceanothus cordulatus
GENERAL DISTRIBUTION:
Whitethorn ceanothus is a characteristic shrub species found in high-elevation brushfields in Baja, California, and mountains of southern California, north to southwest Oregon, and east to the western edge of Nevada [23,27,36,56]. Plants Database provides a distributional map of whitethorn ceanothus.

ECOSYSTEMS [33]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES23 Fir-spruce
FRES26 Lodgepole pine
FRES34 Chaparral-mountain shrub
FRES35 Pinyon-juniper
FRES44 Alpine

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

BLM PHYSIOGRAPHIC REGIONS [8]:
3 Southern Pacific Border
4 Sierra Mountains

KUCHLER [62] PLANT ASSOCIATIONS:
K005 Mixed conifer forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K010 Ponderosa shrub forest
K018 Pine-Douglas-fir forest
K020 Spruce-fir-Douglas-fir forest
K023 Juniper-pinyon woodland
K029 California mixed evergreen forest
K034 Montane chaparral

SAF COVER TYPES [29]:
207 Red fir
210 Interior Douglas-fir
211 White fir
218 Lodgepole pine
219 Limber pine
234 Douglas-fir-tanoak-Pacific madrone
237 Interior ponderosa pine
239 Pinyon-juniper
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
249 Canyon live oak
256 California mixed subalpine

SRM (RANGELAND) COVER TYPES [87]:
209 Montane shrubland
419 Bittercherry
420 Snowbrush

HABITAT TYPES AND PLANT COMMUNITIES:
Whitethorn ceanothus has a narrow range of environments in which it thrives. It is most commonly found within sclerophyllous shrub communities in both conifer and hardwood forests of the coast and interior ranges of the Sierra Nevada [13,31]. In addition to the previously listed habitats, whitethorn ceanothus also occurs in the understory of giant sequoia (Sequoiadendron giganteum) communities/groves because of its ability to tolerate low light levels [17,45,104].

Whitethorn ceanothus is not an indicator species for any specific habitat types, but is listed as a "moderate site indicator" in forested communities of the upper montane in the southern and central Sierra Nevada [79].

Other overstory species commonly associated with whitethorn ceanothus are California black oak (Quercus kelloggii), bitter cherry (Prunus emarginata), and Brewer oak (Q. garryana var. breweri).

Common understory associates of whitethorn ceanothus include: huckleberry oak (Q. vacciniifolia), bush chinquapin (Chrysolepis sempervirens), pinemat manzanita (Arctostaphylos nevadensis), greenleaf manzanita (A. patula), creeping snowberry (Symphoricarpos mollis), redflower currant (Ribes sanguineum), Sierra gooseberry (R. roezlii), California hazel (Corylus cornuta var. californica), deerbrush (Ceanothus integerrimus), prostrate ceanothus (C. prostratus), snowbrush ceanothus (C. velutinus), Sierra mountain misery (Chamaebatia foliolosa), salal (Gaultheria shallon), Pacific rhododendron (Rhododendron macrophyllum), and Saskatoon serviceberry (Amelanchier alnifolia) [21,24,52,60,63,69,98].


 

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Ceanothus cordulatus

 

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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 [25,47,52,74].

Whitethorn ceanothus is a native, evergreen shrub. It has a low-lying, flat-topped growth habit, especially at higher elevations, and generally reaches heights between 2 and 5 feet (0.6-1.5 m). Whitethorn ceanothus can form a continuous ground cover up to 12 feet (3.7 m) in diameter. The many short, rigid, intricately-branched stalks end in sharp spines. The numerous leaves are alternate and distinctly 3-ribbed from the base. They are small, elliptic to egg-shaped, and blunt at the tip. The leaf margins are entire or finely toothed. Flowers are in dense clusters borne on a panicle-like inflorescence. The fruit is a capsule that is slightly crested or horned and somewhat sticky at maturity. It is small, tipped by a threadlike 3-forked appendage, and 3-lobed. Each lobe has a ridge on its back and is 3-celled with each cell bearing 1 hard, round seed or nutlet. [13,25,26,47,73,74, 78,98]

Whitethorn ceanothus has high nitrogen concentrations in its foliage and soil beneath the plant due to the nitrogen-fixing bacteria Frankia spp., suggesting that it may enhance nitrogen availability for the surrounding area [75,76].

RAUNKIAER [83] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Whitethorn ceanothus regenerates by seed and vegetative means. Germination from seeds stored in the soil is the primary mode of reproduction for whitethorn ceanothus and generally takes place after fire [40,54]. Prolific sprouting occurs from a lignotuber when damage is done to the top of the plant [40,50]. Whitethorn ceanothus is an obligate sprouter in moderate fuel-consumption burns and a facultative seeder following high fuel-consumption burns [53].

Pollination: Whitethorn ceanothus is an insect-pollinated species [67].

Breeding system: Outcrossing is the most common breeding system in Ceanothus spp., with limited autogamy [67].

Seed production: Ceanothus spp. are prolific seed producers. Thousands of viable seeds can be produced per hectare and remain dormant in the soil and duff layer until disturbance stimulates germination [38,48]. At approximately 4 years of age, whitethorn ceanothus is capable of seed production and reaches a maximum seed load of approximately 4,500 seeds per plant by the age of 20 to 25. Seed production remains high until the plant is senescent, around 40 years of age, or where the plants are inhibited by shade [18].

Seed dispersal: Whitethorn ceanothus seeds are autochorus, meaning that the main mode of seed dispersal is by the plant itself [55]. The seeds are small and tend to stay where they fall, unless they are carried off by birds, rodents, or ants [22,67]. Ceanothus seeds can also be forcibly ejected from the seed pods when they ripen [22,55].

Seed banking: Because of the large number of seeds produced by whitethorn ceanothus, substantial seed reservoirs exist in the soil and duff layer. The number of seeds in the soil can be expected to vary from year to year depending on the amount of seeds produced and seeds utilized by consumers [22]. Keeley [55] notes a relationship between the seed bank size and stand age where whitethorn ceanothus is found; the older the stand the larger the seed bank will be. Seed quantities of whitethorn ceanothus combined with littleleaf ceanothus (Ceanothus parvifolius) in over-mature mixed coniferous forests can number up to 1.9 million/acre [68].

Germination: Whitethorn ceanothus is heavily dependent upon fire for germination [11]. The seeds of whitethorn ceanothus are refractory: they require a heat stimulus alone or in conjunction with other conditions such as cold stratification [55]. Fire, or similar heat treatment, is necessary to crack the seed coat to allow for absorption of water and subsequent germination. Maximum germination occurs with the sequence of fire and a natural stratification period of 10 to 16 weeks [10,17,38,55,80,82]. Whitethorn ceanothus was observed to be germinating profusely on logged areas, perhaps because of heat from insolation [86].

Fire aids in the germination of seeds but can also destroy them. High intensity or high frequency fires can be detrimental to the seed bank and can result in lower rates of germination [22,68,80,82]. Overall, fire appears to be beneficial in promoting seed numbers, germination, and seedling population [52,68].

The initiation of germination depends on the following: how long the seeds had been dormant; length of time since the last soil disturbance; heat of previous fires; amount of moisture in the soil; persistence of the winter snow pack; soil temperatures; sun exposure; and how deeply the seed was buried [18].

Seedling establishment/growth: Gratkowski [37] notes that whitethorn ceanothus seedling establishment is much higher in burned areas than in nonburned areas, where it is almost nonexistent. Biswell [11] states that seedlings are seldom seen except where there has been fire. Nevertheless, the mortality of Ceanothus spp. seedlings is high during the first 2 postfire years [22]. Environmental factors that affect the growth and survival of these seedlings include insects and other animal browsing, damping-off fungus, soil moisture, exposure, and soil temperature. The first 2 weeks after emergence are the most critical for survival [18].

Asexual regeneration: Layering is common in whitethorn ceanothus at higher elevations where the weight of snow forces branches and stems down to the ground [61]. Stump-sprouting from a lignotuber is also common after the top of the shrub is damaged [40,50].

SITE CHARACTERISTICS:
Whitethorn ceanothus grows best in open situations [10,17]. Dry, open flats, pine forests, rocky ridges, and washes that have well-drained soils are common sites where whitethorn ceanothus is found [23,47,52,74,98]. Areas that have been burned or logged are very favorable for populations of whitethorn ceanothus [2,27,71]. It can also survive and thrives in the shade of coniferous timber [98].

Whitethorn ceanothus is known to occur between 2,100 to 11,000 feet (650-3,350 m) in elevation [7,14,23,47,52,74, 102]

SUCCESSIONAL STATUS:
Whitethorn ceanothus can be classified as both an early- and late-successional species. It is considered a pioneer species most commonly associated with early successional stages because of its ability to germinate from seed and sprout after disturbance [15,81]. Since whitethorn ceanothus is one of the 1st plants to become established on denuded soils, it has immediate value in soil protection and later serves as a nurse crop for coniferous species [98]. It is possible that whitethorn ceanothus plays an important role in succession by providing a more favorable microclimate of nutrient rich microsites in otherwise harsh growing conditions [76].

In normal seral conditions whitethorn ceanothus is overtopped and killed by the conifer species that become established in their shade. The brushfields can, however, significantly hinder conifer regeneration and slow the rate of forest succession. If there is repeated fire, whitethorn ceanothus brushfields can become semipermanent communities [31]. Where fire is lacking, it does not establish and populations decrease, and in some cases are eliminated, in the shade of dense tree canopies [10].

A study by Fernau and others [30] along the west face of the central Sierra Nevada classified whitethorn ceanothus as a late-successional species, with populations peaking 26-32 years after disturbance. Findings published by Minnich [70] also provide evidence that it is capable of existing in later successional stages. He states that chronosequences of postsuccession show the establishment of seedlings of whitethorn ceanothus without fire, apparently from seed caches.

SEASONAL DEVELOPMENT:
Seasonal development of whitethorn ceanothus begins with emergence and extends through a growing season ranging from 75 to 95 days depending on emergence date and elevation at which it is found. Germination of seedlings begins in late May and June, and they are usually well established by 4 weeks of age. The seedlings were observed to grow at a rate of 0.2 inches (0.5 cm) per week until late August when they began to go dormant. After the 1st growing season whitethorn ceanothus grow very rapidly. In mature whitethorn ceanothus new growth and flower bud swelling occur at the same time, ranging from late May to mid-June depending on elevation. Blossoming is expected about 10 days later and continues for approximately 23 days [18].

Whitethorn ceanothus flowers between May and August dependent on where location [23,52,74,78]. Seed maturity is reached in the late summer, generally in August or September.


FIRE ECOLOGY

SPECIES: Ceanothus cordulatus
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Whitethorn ceanothus is highly dependent on fire [11,14]. After fire it regenerates by stump-sprouting or from seed depending on the severity of the burn. It is characterized specifically as an obligate sprouter in moderate fuel-consumption burns and a facultative seeder following high fuel-consumption fires [25,54]. The seeds of whitethorn ceanothus are refractory: requiring heat alone or in conjunction with other conditions such as cold stratification [55]. Heat from a burn initiates germination by cracking the seed coat to allow for absorption of water [10,17,38,80,82].

Fire regimes: Whitethorn ceanothus occurs in a mixed fire regime. The presettlement fire regime for communities where whitethorn ceanothus occurs consisted of frequent, low-severity burns and larger stand-replacing fires. Postsettlement, this trend has changed and produced more lethal fires with higher severities and longer intervals [16]. The fire return intervals in the montane chaparral, where whitethorn ceanothus most often occurs, are "probably quite variable" due to the influence of poor growing conditions [88]. Fire in the subalpine forest habitats where whitethorn ceanothus occurs is infrequent due to limited productivity and fuel accumulation in association with short growing season and heavy snow cover [70].

The following table provides fire return intervals for plant communities and ecosystems where whitethorn ceanothus is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

Community or ecosystem Dominant species Fire return interval range (years)
California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100
pinyon-juniper Pinus-Juniperus spp. <35 [77]
Sierra lodgepole pine* Pinus contorta var. murrayana 35-200
Jeffrey pine Pinus jeffreyi 5-30
Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 [4]
interior ponderosa pine* Pinus ponderosa var. scopulorum 2-30 [4,5,64]
coastal Douglas-fir* Pseudotsuga menziesii var. menziesii 40-240 [4,72,84]
California mixed evergreen Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii <35
canyon live oak Quercus chrysolepis <35 to 200 [4]
California black oak Quercus kelloggii 5-30 [77]
*fire return interval varies widely; trends in variation are noted in the species review

POSTFIRE REGENERATION STRATEGY [93]:
Tall shrub, adventitious bud/root crown
Ground residual colonizer (on-site, initial community)

FIRE EFFECTS

SPECIES: Ceanothus cordulatus
IMMEDIATE FIRE EFFECT ON PLANT:
Fire top-kills or kills whitethorn ceanothus, depending on severity [76]. It is resistant to fire-induced mortality because of the deeply buried meristematic tissues [54].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
A study in mixed conifer ecosystems in the Sierra Nevada, in which whitethorn ceanothus occurred, showed that if duff consumption was 64% or less, few whitethorn ceanothus shrubs were killed and few seeds were stimulated to germinate. The threshold heat load necessary to scarify high numbers of seed stored in the soil bank was very close to the threshold that can kill below ground meristematic tissues of resprouting shrubs. At consumption levels greater than 64% the number of seedlings increased exponentially while survival of resprouters decreased from 94 to 50%. If duff consumption was greater than 94% both are reduced and the heat loads are severe enough to cause a decline in the number of seeds in the soil which survive and successfully germinate [54]. A later study by Kauffman and Martin [53] done in the same mixed conifer ecosystems of the Sierra Nevada , shows that whitethorn ceanothus was more resistant to fire-induced mortality than other species found in association, such as California black oak and tanoak (Lithocarpus densiflorus). Although there were few or relatively no shrubs present on plots before burning, numbers and frequency of whitethorn ceanothus increased after burning [46,59].

PLANT RESPONSE TO FIRE:
Whitethorn ceanothus responds favorably to fire. Many soil-stored seeds germinate after fire breaks seed dormancy; seedlings can account for more than half of the plants present in burn areas [34,37]. Whitethorn ceanothus also stump-sprouts after fire [25]. Burned areas stimulate nitrification through the perpetuation of nitrogen-fixing shrubs such as whitethorn ceanothus, accelerating the recycling of nutrients stored in living and dead plants [14].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Dormant whitethorn ceanothus seeds in the soil can withstand temperatures up to 220 °F (105 ºC). This ability to withstand high soil temperatures is an important survival tool and permits the seeds to endure long heat exposure during wildland fires and then germinate and occupy burned areas. Seeds exposed to lethal temperature regimes created by burning heavy accumulations of slash or decaying logs are destroyed [38].

Fire removes most of the above ground whitethorn ceanothus biomass, leaving nitrogen-rich, bare mineral soil. Following fire, patches of whitethorn ceanothus maintain higher amounts of available nitrogen for up to 6 months on both low- and high-intensity burn plots. If these areas continue to have more available nitrogen than other patch types, this long-term effect may enhance tree seedling growth [76].

The Research Project Summary Plant response to prescribed burning with varying season, weather, and fuel moisture in mixed-conifer forests of California provides information on prescribed fire and postfire response of many plant community species including whitethorn ceanothus.

FIRE MANAGEMENT CONSIDERATIONS:
Prescribed burning of whitethorn ceanothus is used in land management to increase browse [27].

The evolution of whitethorn ceanothus and fire together suggests that the use of prescribed burning is a viable management option in areas to promote whitethorn ceanothus. The use of fire to promote whitethorn ceanothus is suitable because of the fire-stimulated germination of seeds and stump-sprouting in moderate fuel consumption burns. The species adds nitrogen to the soil, possibly enhancing conifer regeneration.

Conversely, overstocked or well-established stands of whitethorn ceanothus add significantly to fuel loading. Extensive stands of whitethorn ceanothus and montane chaparral patches exhibited the greatest mean fire spread/minute and the greatest flame lengths compared to forested types [101]. Snyder [91] notes that during succession, brush cover is still significant even as the basal area of trees doubles. If these populations persist as noted they could act as ladder fuels, increasing the possibility of a stand replacing fire.

Fuel loading becomes an important factor to recognize when using prescribed fire as a land management tool. The changing fire regime in communities where whitethorn ceanothus becomes established, to infrequent, high-severity fires [16], suggests that using fire to manage whitethorn ceanothus should be done with caution.


MANAGEMENT CONSIDERATIONS

SPECIES: Ceanothus cordulatus
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Whitethorn ceanothus is an important browse producer for wildlife species [10]. It is a major component of the diet of mule deer in the summer ranges of the Sierra Nevada and is also used by cattle on summer grazing allotments [9,17,27,49,57,58,85]. Gordon [34] suggests that the spine-tipped branchlets that develop after establishment may make the shrub less desirable to deer as a browse component. Whitethorn ceanothus is also browsed by bighorn sheep, goats, and domestic sheep [22,25]. Long-eared chipmunks collect and cache the seeds of whitethorn ceanothus [41,97]. High amounts of whitethorn ceanothus are eaten by pocket gophers in California [96]. The fruit of the shrub is eaten by birds and small mammals [25]. Whitethorn ceanothus also provides forage and nesting habitat for the dusky woodrat and part of the diet of the mountain pocket gopher [105]. Whitethorn ceanothus is a host plant for the Great Basin tent caterpillar [20].

Palatability/nutritional value: Whitethorn ceanothus is an important browse species for deer because of its high crude protein content and palatability [9,22,25,49]. It does not, however, provide sufficient nutrients for optimum growth and development in mule deer and is more valuable when consumed along with other species [58]. It is considered as a poor to fair browse species for livestock such as cattle, domestic sheep, and goats [22,25,73].

Cover value: No information is available on this topic.

VALUE FOR REHABILITATION OF DISTURBED SITES:
Whitethorn ceanothus has considerable value when used for site amelioration. It is used for erosion control on slopes, terraces, and steep banks [78,89,90]. Whitethorn ceanothus is very important for rehabilitation because of nitrogen-fixing and other soil-building attributes. It is tolerant of severe sites and can play a soil building role on these sites [22,49]. It has successfully been used for these purposes in the Lake Tahoe Basin and Lost Canyon, California [15,19,89,90].

A list of available/potential seed sources can be acquired in the USDA, NRCS Commercial Sources of Conservation Plant Material, available online [99].

OTHER USES:
No information is available on this topic.

OTHER MANAGEMENT CONSIDERATIONS:
There a few main points to consider when managing whitethorn ceanothus. It increases with disturbance and forms very dense, difficult to penetrate brushfields. It is regarded as a very strong competitor that can hinder tree seedling growth and subsequent conifer regeneration [22,57,94]. Whitethorn ceanothus may compete with tree seedlings, but it also provides ongoing nitrogen that may improve the site for conifer establishment and growth after establishment. Soil nitrogen, however, probably becomes most important for tree seedlings after they have cleared the initial hurdle of establishment [49,76]. In the past, herbicide treatments and allowing livestock grazing in whitethorn ceanothus brushfields were used as management options to open the dense shrub canopy. This increased the amount of light reaching the young conifers in the understory and decreased brush competition for soil moisture and nutrients. Initial applications of herbicide resulted in 100% top kill and 20% whitethorn ceanothus mortality with plant mortality reaching 90% after the 3rd application [35,39,40,43,92,103]. A study done by Hobbs and Radosevich [48] concluded that the control of evergreen woody species such as whitethorn ceanothus in young conifer plantations improved tree seedling growth and in some cases survival.

Conversely, whitethorn ceanothus is valuable as a browse species and if populations decrease, the amount of available forage for wildlife and livestock is also decreased [57]. Prescribed burns are often used in land management to encourage the establishment of whitethorn ceanothus because of its importance as a browse species [27]. It is also beneficial as a nurse plant for conifer and Sierra gooseberry seedlings, providing shade and available nitrogen [81].

Tappeiner [95] reported the average size of open-grown whitethorn ceanothus plants on ungrazed, 9-year-old tree plantations in the Sierra Nevada and Klamath Mountains. Average crown diameter was 5.7 feet (1.7 m) and average height was 3.2 feet (.98 m). This information may be useful in predicting shrub development.


Ceanothus cordulatus: REFERENCES


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27. Eberlein, Gary P. 1982. Estimating growth of young mountain whitethorn shrubs. PSW-357. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 3 p. [6351]
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