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SPECIES:  Frangula californica
California buckthorn. Image ©2012 Jean Pawek, used with permission.

 


Introductory


AUTHORSHIP AND CITATION: McMurray, Nancy E. 1990. Frangula californica. 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.fed.us/database/feis/plants/shrub/fracal []. Updates: On 10 July 2018, the common name of this species was changed from: California coffeeberry to: California buckthorn. Images were also added. ABBREVIATION: FRACAL NRCS PLANT CODE [79]: FRCA12 COMMON NAMES: California buckthorn California coffeeberry California false buckthorn hoary coffeeberry TAXONOMY: The scientific name of California buckthorn is Frangula californica (Eschsch.) Gray (Rhamnaceae). There are 6 subspecies [37,79,81]: Frangula californica subsp. californica Frangula californica subsp. crassifolia (Jep.) Kartesz & Gandhi Frangula californica subsp. cuspidata (Greene) Kartesz & Gandhi Frangula californica subsp. occidentalis (J. Howell) Kartesz & Gandhi Frangula californica subsp. tomentella (Benth.) Kartesz & Gandhi, hoary coffeeberry Frangula californica subsp. ursina (Greene) Kartesz & Gandhi SYNONYMS: Rhamnus californica Esch. Rhamnus californica subsp. californica Rhamnus californica subsp. occidentalis (J. Howell) C. Wolf Rhamnus tomentella Benth. Rhamnus tomentella Benth. subsp. crassifolia (Jeps.) J.S. Sawyer Rhamnus tomentella Benth. subsp. cuspidata (Greene) J.S. Sawyer Rhamnus tomentella Benth. subsp. ursina (Greene) J.S. Sawyer [76] LIFE FORM: Shrub FEDERAL LEGAL STATUS: No special status OTHER STATUS: NO-ENTRY


DISTRIBUTION AND OCCURRENCE

SPECIES: Frangula californica
GENERAL DISTRIBUTION: California buckthorn ranges from extreme southwestern Oregon southward along the coast and Coast Ranges to southwestern California 976].  It is cultivated in Hawaii [80].
Distribution of California buckthorn. Map courtesy of USDA, NRCS. 2018. The PLANTS Database. National Plant Data Team, Greensboro, NC [2018, July 10] [79].
ECOSYSTEMS: 
   FRES20  Douglas-fir
   FRES21  Ponderosa pine
   FRES23  Fir-spruce
   FRES27  Redwood
   FRES28  Western hardwoods
   FRES34  Chaparral - mountain shrub


STATES: 
     CA  HI  OR


BLM PHYSIOGRAPHIC REGIONS: 
    1  Northern Pacific Border
    3  Southern Pacific Border


KUCHLER PLANT ASSOCIATIONS: 
   K005  Mixed conifer forest
   K006  Redwood forest
   K007  Red fir forest
   K009  Pine - cypress forest
   K012  Douglas-fir forest
   K029  California mixed evergreen forest
   K030  California oakwoods
   K033  Chaparral
   K035  Coastal sagebrush
   K036  Mosaic of K030 and K035


SAF COVER TYPES: 
   207  Red fir
   229  Pacific Douglas-fir
   231  Port-Orford-cedar
   232  Redwood
   234  Douglas-fir - tanoak - Pacific madrone
   248  Knobcone pine
   249  Canyon live oak
   250  Blue oak - gray pine
   255  California coast live oak


SRM (RANGELAND) COVER TYPES: 
   201  Blue oak woodland
   202  Coast live oak woodland
   203  Riparian woodland
   204  North coastal shrub
   205  Coastal sage shrub
   207  Scrub oak mixed chaparral


HABITAT TYPES AND PLANT COMMUNITIES: 
California buckthorn is a shrub component of chaparral, woodland, and
forest communities throughout its distribution [4,8,52,55,56].  It has
not been used as an indicator species in published classification
schemes for California.  In the Siskiyou Mountains of southwestern
Oregon and extreme northern California, Atzet and Wheeler [1] describe a
tanoak/California buckthorn (Lithocarpus densiflorus/Rhamnus
californica) plant association on ultrabasic parent materials.  Although
the tanoak series typically occurs on deep, fertile soils in the
Siskiyou Mountain province, this association is the ultrabasic version
of a tanoak climax.  Dominance of coffeeberry apparently indicates a
soil imbalance [1].

In southern and central California, California buckthorn is a frequent
member of coastal chaparral and sage scrub [13,18] and is most commonly
associated with relatively mesic scrub oak (Quercus spp.) chaparral
[30].  It also becomes locally abundant on cool, fog-dominated sites
along the central Coast Range where it occurs beneath mixed-hardwood
forests dominated by coast live oak (Quercus agrifolia) [15,54,65,70].
Although never very abundant, California buckthorn is often associated
with woodland and forest mosaics throughout southern and central
California.  Within oak woodlands, knobcone pine (Pinus attenuata), and
coastal forests, it occurs both as a scattered understory shrub and as a
component of intermixed stands of "woodland chaparral" [35,30,67,68].
Some common associates are chaparral whitethorn (Ceanothus leucodermis),
toyon (Heteromeles arbutifolia), skunkbush sumac (Rhus trilobata),
redberry (Rhamnus crocea), hollyleaf redberry (R. crocea var.
ilicifolia), and poison-oak (Toxicodendron diversilobum).

In northern California and southwestern Oregon, California buckthorn
occurs in mixed evergreen, red fir (Abies magnifica var. shastensis),
and redwood (Sequoia sempervirens) forests [49,52,65].  On
mixed-evergreen sites with ultrabasic parent materials, the
sclerophyllous subcanopy is largely replaced by a shrub layer dominated
by California buckthorn (ssp. occidentalis) and evergreen huckleberry
(Vaccinium ovatum) [1,64,65].  The open, conifer overstory is usually
composed of Port-Orford cedar (Chamaecyparis lawsoniana), knobcone pine,
sugar pine (Pinus lambertiana), and Douglas-fir (Pseudotsuga menziesii)
with a sparse subcanopy of tanoak, huckleberry oak (Quercus
vaccinifolia), and California laurel (Umbellularia californica)
[1,2,3,65].

California buckthorn is also a characteristic shrub within extensive,
evergreen brushfields in the Siskiyou Mountain province [21,24,25].
Common brushfield associates include whiteleaf manzanita (Arctostaphylos
viscida), greenleaf manzanita (A. patula), hoary manzanita (A.
canescens), wedgeleaf ceanothus (Ceanothus cuneatus), and deer brush (C.
integerrimus) [21,25].

MANAGEMENT CONSIDERATIONS

SPECIES: Frangula californica
IMPORTANCE TO LIVESTOCK AND WILDLIFE: California buckthorn is distributed throughout much of California and is generally considered a staple browse of both big game and livestock [12].  Browse is more greater important to sheep, goats, and deer than to cattle [17,63,74].  Use typically occurs in the fall when more palatable herbaceous plants are cured [50].  California buckthorn is an important mule deer browse on winter ranges in portions of California [17].  The fruits of California buckthorn are extensively utilized by numerous wildlife species, particularly birds [12,17].  In many areas, the berries are often the only abundant "juicy" fruit available in the fall [75].  Berries are readily eaten by band-tailed pigeons, black-tailed deer, and black bears [12,74].  Woodrats eat limited quantities of the seeds [33]. PALATABILITY: The palatability of California buckthorn browse depends upon stem age and community associates [17,63].  Cattle utilization is usually limited, particularly in communities where California buckthorn has a scattered distribution.  However, where plants are locally abundant, the current annual growth is often heavily utilized [63].  postfire sprouts are highly preferred by livestock and big game [17,27,63]. Browse ratings for California buckthorn are presented below [63]:     sheep                good - poor     goats                good - poor     deer                 good - poor     cattle               fair - poor     horses               useless NUTRITIONAL VALUE: Nutritionally, California buckthorn is a satisfactory food source for livestock and big game [63].  Although foliar protein content is never particularly high, California buckthorn is important because it remains succulent throughout the year [23].  Nutritional studies of the foliage indicate that protein content of mature leaves is low (7.5 percent) from November through March.  Maximum values are reached between April and August when the protein content of newly developed foliage reaches an average of 19 percent [63].  Nutrient content of the leaves compares favorably with common shrub associates such as wedgeleaf ceanothus and chaparral whitethorn [23].  Crude fiber values show little seasonal variation, ranging from 13.5 percent in newly initiated leaves to 15 percent in mature foliage [63]. COVER VALUE: The cover value of California buckthorn has not been documented. Tanoak/coffeeberry associations in southern Oregon and northern California are characterized by a diverse vertical structure, supplying nesting cover and perching sites for numerous bird species [1]. VALUE FOR REHABILITATION OF DISTURBED SITES: California buckthorn is useful for erosion control on dry, steep hillsides [19].  One- to two-year-old transplants produce substantial seed crops and large numbers of volunteers occur on sites receiving additional irrigation.  On suitable sites, transplants may grow 8 to 12 feet (2.4-3.7 m) tall with comparable spreads within approximately 20 years [19].  Since the berries are highly preferred by a variety of bird species, California buckthorn is a good selection for wildlife plantings [34]. Plants are easily propagated from seed sown in nursery beds using either fresh, unstratified seed in the fall or stratified seed in the spring [34,74].  Seed should be collected in the fall approximately 2 weeks before it is fully ripe, and the pulp removed prior to sowing [34]. When stored in sealed containers at 41 degrees F (5 degrees C), buckthorn (Rhamnus spp.) seed remains viable for several years [34]. Propagation from stem cuttings is difficult but can be achieved using soft or hardwood cuttings made at the nodes [34,43,74]. OTHER USES AND VALUES: California buckthorn is frequently cultivated for ornamental purposes since the shiny, colorful berries contrast nicely with the light green foliage [19,60,63].  The berries are sweet and edible and were gathered historically by West Coast Indian tribes for culinary as well as medicinal purposes [12].  Although the berries superficially resemble the commercial coffee bean, attempts at using California buckthorn as a coffee substitute have not been successful.  The bark was once exported for use as a laxative [14]. OTHER MANAGEMENT CONSIDERATIONS: Planting programs:  Some species belonging to the buckthorn (Rhamnus) genus serve as alternate hosts for the oat rust, Puccinia coronata [34]. California buckthorn, however, is a secondary host for the rust of velvet grass (Holcus spp.) and is not a concern when planted near fields of cultivated oats [74]. Herbicides:  California buckthorn is sensitive to almost all forestry-registered herbicides [11].  If sprouts are treated following burning, plants are usually killed by retreatment [7,9,26,69].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Frangula californica
GENERAL BOTANICAL CHARACTERISTICS: California buckthorn is a native, broad-leaved, sclerophyllous shrub [13,63,74].  Growth habit varies according to subspecies with plants ranging from low, spreading shrubs to upright, arborescent individuals [52].  On favorable sites along the coast, California buckthorn occasionally grows as a small tree, reaching heights of approximately 20 feet (6.1 m) [45].  More often, however, it grows as a 4 to 6 foot (1.2-1.8 m) tall shrub [63].  Bark of young twigs is usually reddish; older branches have gray, brown, or reddish bark [12,52].  The small, pinnately veined, evergreen leaves are commonly dark green above and paler beneath, and are arranged alternately on the stem [53].  Leaf margins are typically inrolled [12,14].  When growing on xeric sites, leaves tend to be small and thick; in moist situations they are relatively large and thin [63].  The inconspicuous, bisexual flowers are green and occur in small, axillary clusters [53].  The fruit is a juicy, berrylike drupe approximately 0.25 inch (7-9 mm) in diameter and may be either green, black, or red in color [34,52].  Berries contain two smooth, nutlike seeds which closely resemble the commercial coffee bean [12,14,34].  Although the root crown may become enlarged in response to repeated postfire sprouting, this structure is not a lignotuber [38,40]. Longevity of California buckthorn is estimated at 100 to 200 years [39]. RAUNKIAER LIFE FORM:       Phanerophyte REGENERATION PROCESSES: California buckthorn regenerates by both sexual and vegetative means. On chaparral sites in southern California, California buckthorn maintains itself primarily through sprouting [41].  Seedling establishment is never very abundant and is restricted to stands of mature chaparral [39,40,41].  Little or no seedling establishment occurs immediately following fire [38,40,41].  However, seemingly different establishment patterns have been observed in other communities [63,66] Vegetative regeneration:  In the absence of fire, many long-lived sprouters within stands of mature chaparral rejuvenate their canopies by continually producing new sprouts from established root crowns [40,42]. Generalized information indicates that California buckthorn may also maintain itself in this manner [40].  Following disturbances such as fire or cutting, California buckthorn sprouts from surviving adventitious buds on the root crown [36,62]. Seed reproduction:  Onset of seed production occurs early in California coffeeberry, usually by 2 to 3 years of age [19].  Seeds are dispersed in the fall [41].  Significant, widespread dispersal of the pea-sized berries occurs through animals, particularly birds [10,41].  Bird harvest of the fruit crop is often so complete that relatively few seeds fall beneath the parent plant.  The seeds of California buckthorn are apparently quite short lived.  When dried at room temperature, viability is retained for no longer than 9 months.  At the time of dispersal, each seed exhibits a chlorophyllous cotyledon, indicating that germination is imminent [41].  If kept too moist prior to germination, seeds are prone to rot [19].  Germination occurs readily under favorable moisture and temperature conditions [34,41].  Keeley [41] recently studied the germination requirements of California buckthorn using seed samples collected in southern California.  After a 1-month stratification at 41 degrees F (5 degrees C), 65 percent of California buckthorn seeds germinated when light incubated at 73 degrees F (23 degrees C) for 3 weeks.  Rate of germination was rapid with more than 75 percent of germination occurring during the first week.  Addition of charate (powdered charred wood) greatly reduced germination under similar conditions (15% germination); in the dark, however, addition of charate stimulated germination relative to the control (90% germination).  In this study, heat treatments generally decreased germination.  Longer heating at low temperatures was more detrimental than short bursts of high temperature [41].  Sampson [62] found that heat treatments produced a slight increase in germination. Keeley [38,39,40,41] reported that buckthorns (Rhamnus spp.) are obligate sprouters after fires in southern California chaparral and included both California buckthorn and redberry (Rhamnus crocea) within this grouping.  Obligate sprouting species are restricted to sprouting following fire and do not establish seedlings in the initial postfire environment.  In fact, seedling establishment of obligate sprouters is always quite limited and follows the generalized scenario presented below [40,41,71,72]:      -- Seedlings are established primarily in mature chaparral         in gaps resulting from the death of senescing, shorter-lived         species.      -- Seedling establishment is often episodic and coincides with         periods of above normal rainfall .      -- Although initial establishment may occur in burned or         unburned stands during very wet years, continued survival is         favored beneath mature stands on sites that are relatively         mesic (north slopes) and which possess a well-developed litter         layer.      -- Long-term survival beneath mature chaparral is rare; seedlings         are stunted and are subjected to heavy browsing by small         mammals.      -- Seedlings are most common in very old stands (60 to 100+         years) where long fire free intervals allow for the build         up of seedling populations. Redberry tends to follow the above pattern [28,29,31,32,42,57], but data is scant concerning the seedling ecology of California buckthorn.  A review of the literature found no data on California buckthorn seedling establishment within southern California chaparral.  On Coast Range sites in northern California, Sampson [62] did not observe any California buckthorn seedlings beneath stands of manzanita-ceanothus (Arctostaphylos spp.-Ceanothus spp.) chaparral.  Seedlings were present on adjacent burns.  Densities equaled 4,400 seedlings/acre (10,872 seedlings/ha) 1 year after fire; 4 years later, there were approximately 2,300 seedlings/acre (5,683 seedlings/ha).  Precipitation was apparently below normal during the first three postfire growing seasons [62]. Pelton [58] found occasional California buckthorn seedlings beneath mixed-hardwood forests in the Santa Cruz Mountains.  Seedlings of var. occidentalis apparently require some shade during the establishment period [19]. SITE CHARACTERISTICS: California buckthorn exhibits a wide ecological amplitude.  Sites include dry flats, moist slopes, ravines, and rocky ridges, usually at elevations below 5,500 feet (1,677 m) [52,63].  Soils are typically dry and well drained [74].  Established plants tolerate full sun to moderate shade [13,74]. In the Siskiyou Mountains, sites supporting tanoak/coffeeberry plant associations occur on flat, lower slope positions at elevations ranging from 1,040 to 3,460 feet (317 to 1,055 m) on southerly aspects; ultrabasic soils reach depths of approximately 30 inches (76 cm). SUCCESSIONAL STATUS: California buckthorn is a long-lived and moderately shade-tolerant shrub that is highly persistent within chaparral, hardwood woodland, and open conifer forests [13,39].  During extended fire free-intervals, California buckthorn is able to outlive, overtop, and shade out many shorter-lived species [71].  As a component of relatively open canopied stands, plants persist until the next fire occurs [36,66], at which time sprouted individuals become part of the initial postfire vegetation [66].  Griffin [75], however, observed many senescing plants of California buckthorn on mixed hardwood sites in the southern Coast Range; according to Griffin, California buckthorn had been the dominant shrub on these sites for quite some time.  Shrubs with bird-dispersed seed, such as California buckthorn, have apparently increased in abundance on relic oak savanna sites in central California [35]. SEASONAL DEVELOPMENT: California buckthorn typically flowers from April to June [34,52]. Fruit ripening commonly occurs from July through November with dispersal taking place during the fall [34,41,74].  Immature fruits are green, turning red or reddish-black when fully ripened [53,63].  Although an evergreen species, California buckthorn produces at least some new leaves each year [23].  Observations on the phenological development of California buckthorn during 1936 for sites in Shasta County, California, are presented below [62]:     Phenological stage                           Date     Leaves half developed                        April 11     Leaves fully developed                       May 16     Leaves fully developed & fruit forming       June 24

FIRE ECOLOGY

SPECIES: Frangula californica
FIRE ECOLOGY OR ADAPTATIONS: Following fires which kill aerial stems, California buckthorn sprouts vigorously from dormant buds located on the root crown [62].  The root crown serves as a source of numerous perennating buds and stored carbohydrates, enabling California buckthorn to rapidly reoccupy the initial postfire environment [41,48]. FIRE REGIMES: 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". POSTFIRE REGENERATION STRATEGY:    Tall shrub, adventitious-bud root crown

FIRE EFFECTS

SPECIES: Frangula californica
IMMEDIATE FIRE EFFECT ON PLANT: California buckthorn is quite resistant to fire mortality [62,66]. Although aerial portions may be top-killed, most plants survive fire [66]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT: NO-ENTRY PLANT RESPONSE TO FIRE: Vigorous sprouting is the primary means by which California buckthorn reestablishes itself in the postfire environment [38,40,41,62]. The degree to which seedlings contribute to its postfire recovery seems to vary with fire intensity, community type, and perhaps geographical location [41,66]. Vegetative regeneration:  California buckthorn sprouts vigorously following fires which kill the aerial stems [6,36,39,63].  Although cover and basal area may be initially reduced following burning [26,47], most plants rapidly regain their prefire size and biomass [59]. Seedling reproduction:  California buckthorn produces short-lived seeds, the majority of which germinate readily under favorable temperature and moisture conditions [34,41,62].  Consequently, postfire establishment may occur through bird dispersal of off-site seed [41,42]. Seed production by residual plants may also be a factor on some sites. Generalized information on obligate sprouters suggests that sprouted plants begin to produce seed crops within 1 to 2 years of burning and that postfire fruit crops are often substantial [40].  Although most seeds are not well adapted to resist fire or for long-term survival in the soil [41], germination in a portion of the seeds may be cued to the postfire environment.  Sampson [62] reported a slight increase in germination when California buckthorn seeds were exposed for 5 minutes to heat treatments of 140 to 180 degrees F (60 to 82 degrees C).  Heat treated samples from both southern California (San Bernardino Co.) and northern California (Mendocino Co.) showed an increase in germination over controls; the greatest increase occurred in the northern California sample [62].  Keeley [41] found that heat treatments generally decreased germination; optimal germination occurred when charred wood was added to dark incubated controls. Limited information presents an unclear pattern of postfire seedling establishment in California buckthorn.  In southern California chaparral, it apparently behaves as an obligate spouter and rarely establishes seedlings in the initial postfire environment [40,41]. Generalized information indicates that seedlings of obligate sprouting species are rarely observed during the first postfire season except during periods of above-normal precipitation [40,73].  Although seedlings may initially establish in fire-created gaps in very wet years, successful establishment seems restricted to mesic sites beneath mature chaparral where litter layers are well developed [40,72]. On chaparral sites in northern California, however, Sampson [62] found seedling densities of 4,400 /acre (10,872 seedlings/ha) on 1-year-old burns.  Although individual seedling survival was not followed, there were 2,300 seedlings/acre (5,683 seedlings/ha) at the end of the fifth postfire season.  Prior to burning, seedlings were not observed beneath adjacent unburned stands consisting of a cover of sprouting manzanita and ceanothus (stand age not given) [62].  California buckthorn seedlings did not establish the first year after a wildfire burned a summit sugar pine forest in the Santa Lucia Range of central California despite its common occurrence in the prefire vegetation [66].  This wildfire was described as intense and burned an extensive area. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE: Response of vegetation to prescribed burning in a Jeffrey pine-California black oak woodland and a deergrass meadow at Cuyamaca State Park, California, provides information on prescribed fire use and postfire response of many mixed-conifer woodland species including California coffeeberry. FIRE MANAGEMENT CONSIDERATIONS: Wildlife management:  Burning initially increases the palatability of California buckthorn browse [27,62,63].  Plants on recently burned sites are higher in crude protein and crude fiber values than plants in unburned stands [62].  Sprouts are generally utilized for up to two postfire growing seasons [7].  On small burns, use of California coffeeberry may be so concentrated that plants are weakened to the point that mortality ensues [27].

REFERENCES

SPECIES: Frangula californica
REFERENCES:  1.  Atzet, Thomas; Wheeler, David L. 1984. Preliminary plant associations of        the Siskiyou Mountain Province. Portland, OR: U.S. Department of        Agriculture, Forest Service, Pacific Northwest Region. 278 p.  [9351]  2.  Atzet, Tom; Wheeler, David; Smith, Brad; [and others]. 1984. The tanoak        series of the Siskiyou Region of southwest Oregon. Forestry Intensified        Research [Oregon State University]. 6(3): 6-7.  [8593]  3.  Atzet, Tom; Wheeler, David; Smith, Brad; [and others]. 1985. The tanoak        series of the Siskiyou region of southwest Oregon (Part 2). Forestry        Intensified Research. 6(4): 7-10.  [8594]  4.  Barbour, Michael G.; Major, Jack, eds. 1977. Terrestrial vegetation of        California. New York: John Wiley & Sons. 1002 p.  [388]  5.  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]  6.  Biswell, H. H. 1959. Prescribed burning and other methods of deer range        improvement in ponderosa pine in California. In: Proceedings, Society of        American Foresters; 1959; San Francisco, CA. Bethesda, MD: Society of        American Foresters: 102-105.  [5269]  7.  Biswell, H. H. 1961. Manipulation of chamise brush for deer range        improvement. California Fish and Game. 47(2): 125-144.  [6366]  8.  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]  9.  Bovey, Rodney W. 1977. Response of selected woody plants in the United        States to herbicides. Agric. Handb. 493. Washington, DC: U.S. Department        of Agriculture, Agricultural Research Service. 101 p.  [8899] 10.  Bullock, Stephen H. 1978. Fruit abundance and distribution in relation        to types of seed dispersal in chaparral. Madrono. 25: 104-105.  [9792] 11.  Burrill, Larry C.; Braunworth, William S., Jr.; William, Ray D.; [and        others], compilers. 1989. Pacific Northwest weed control handbook.        Corvallis, OR: Oregon State University, Extension Service, Agricultural        Communications. 276 p.  [6235] 12.  Conrad, C. Eugene. 1987. Common shrubs of chaparral and associated        ecosystems of southern California. Gen. Tech. Rep. PSW-99. Berkeley, CA:        U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest        and Range Experiment Station. 86 p.  [4209] 13.  Cooper, W. S. 1922. The broad-sclerophyll vegetation of California.        Publ. No. 319. Washington, DC: The Carnegie Institution of Washington.        145 p.  [6716] 14.  Dale, Nancy. 1986. Flowering plants: The Santa Monica Mountains, coastal        and chaparral regions of southern California. Santa Barbara, CA: Capra        Press. In coooperation with: The California Native Plant Society. 239 p.        [7605] 15.  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] 16.  Detling, LeRoy E. 1961. The chaparral formation of southwestern Oregon,        with considerations of its postglacial history. Ecology. 42(2): 348-357.        [6360] 17.  Dayton, William A. 1931. Important western browse plants. Misc. Publ.        101. Washington, DC: U.S. Department of Agriculture. 214 p.  [768] 18.  Dunn, Paul H.; Barro, Susan C.; Wells, Wade G., II; [and others]. 1988.        The San Dimas Experimental Forest: 50 years of research. Gen. Tech. Rep.        PSW-104. Berkeley, CA: U.S. Department of Agriculture, Forest Service,        Pacific Southwest Forest and Range Experiment Station. 49 p.  [8400] 19.  Everett, Percy C. 1957. A summary of the culture of California plants at        the Rancho Santa Ana Botanic Garden 1927-1950. Claremont, CA: The Rancho        Santa Ana Botanic Garden. 223 p.  [7191] 20.  Eyre, F. H., ed. 1980. Forest cover types of the United States and        Canada. Washington, DC: Society of American Foresters. 148 p.  [905] 21.  Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon        and Washington. Gen. Tech. Rep. PNW-8. 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