Index of Species Information

SPECIES:  Adenostoma fasciculatum

Introductory

SPECIES: Adenostoma fasciculatum
AUTHORSHIP AND CITATION : McMurray, Nancy E. 1990. Adenostoma fasciculatum. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].
ABBREVIATION : ADEFAS SYNONYMS : NO-ENTRY SCS PLANT CODE : ADFA ADFAF ADFAO COMMON NAMES : chamise greasewood chamise chamiso TAXONOMY : The currently accepted scientific name of chamise is Adenostoma fasciculatum Hook and Arn. (Rosaceae). Chamise and red shank (A. sparsifolium) are the only members of this genus [38,91]. Chamise has two recognized varieties which are differentiated on the basis of leaf size and shape [64,91]: A. f. var. fasciculatum and var. A. f. var. obtusifolium S. Watson [18,91]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY


DISTRIBUTION AND OCCURRENCE

SPECIES: Adenostoma fasciculatum
GENERAL DISTRIBUTION : Chamise is the most characteristic and widely distributed chaparral species in California [25,38,121]. It is most extensively distributed in the southern Coast Ranges [20,26,48], but occurs in the Coast, Transverse and Peninsular ranges from Mendocino County to Baja California [20,26,48]. It also occurs in the Sierra Nevada foothills [121] and on the Channel islands [26]. Adenostoma fasciculatum variety obtusifolium is restricted to southwestern San Diego County and Baja California [26,92]. ECOSYSTEMS : FRES21 Ponderosa pine FRES28 Western hardwoods FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper STATES : CA MEXICO BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 3 Southern Pacific Border 4 Sierra Nevada KUCHLER PLANT ASSOCIATIONS : K009 Pine - cypress forest K023 Juniper - pinyon woodland K030 California oakwoods K033 Chaparral K035 Coastal sagebrush K036 Mosaic of K030 and K035 SAF COVER TYPES : 239 Pinyon - juniper 245 Pacific ponderosa pine 246 California black oak 248 Knobcone pine 249 Canyon live oak 250 Blue oak - Digger pine 255 California coast live oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Chamise is a shrub component of chaparral, woodland, and forest communities throughout much of California [13,51]. Within chaparral communities, chamise typically dominates the shrub cover on the hottest and driest sites [102]. As available moisture increases, it codominates with manzanita (Arctostaphylos spp.) and ceanothus (Ceanothus spp.) species [42,51]. Chamise is an understory shrub in dry coniferous woodlands dominated by Parry pinyon (Pinus quadrifolia), knobcone pine (P. attenuata), or gray pine (P. sabiniana). Less commonly, chamise occurs beneath scrubby "forest" communities dominated by either Torrey pine (P. torreyana), knobcone pine, Piute cypress (Cupressus arizonica ssp. nevadensis), Cuyamaca cypress (C. a. var. stephensonii), or Tecate cypress (C. forbesii) [51]. It is also present in the understory of maritime Coast Range ponderosa pine (Pinus ponderosa) forests [51]. Common associates within chamise chaparral include [13]: northern Coast Range: hoary manzanita (Arctostaphylos canescens), Parry manzanita (A. manzanita), wedgeleaf ceanothus (Ceanothus cuneatus), wavyleaf ceanothus (C. foliosus), and leather oak (Quercus durata). southern Coast Range: oaks (Quercus spp.), ceanothus (Ceanothus spp.), manzanitas (Arctostaphylos spp.), buckthorns (Rhamnus spp.), sumacs (Rhus and Malosma spp.), California sagebrush (Artemisia californica), California buckwheat (Eriogonum fasciculatum), and sage (Salvia spp.). interior: whiteleaf manzanita (Arctostaphylos viscida), Parry manzanita, wedgeleaf ceanothus, Lemmon ceanothus (C. lemmonnii), chaparral whitethorn, toyon (Heteromeles arbutifolia), buckthorns, poison-oak (Toxicodendron diversilobum), and yerba santa (Eriodictyon californicum). southern California: bigberry manzanita (A. glauca), Mexican manzanita (A. pungens), pink-bracted manzanita (A. pringlei var. drupacea), hoaryleaf ceanothus (C. crassifolius), and desert ceanothus (C. greggi var. perplexans). Published classifications listing chamise as a dominant or indicator species include: The chaparral vegetation of Santa Cruz Island, California [11] Vegetation and floristics of Pinnacles National Monument [36] Vegetation types of the San Gabriel Mountains [41] Preliminary descriptions of the terrestrial natural communities of California [51] Vegetation types of the San Bernardino Mountains [53] A vegetation classification system applied to southern California [102]

MANAGEMENT CONSIDERATIONS

SPECIES: Adenostoma fasciculatum
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Chamise is important forage more because of its abundance and widespread distribution than its palatability [121]. It furnishes a large quantity of low to medium quality browse for both livestock and big game [121]. Chamise is a staple deer food (by volume) throughout much of California [76,121]. Deer use is often year-round, particularly in northern California [9,121], but is most concentrated during the summer and fall [9,50,76,121]. Limited livestock use occurs, primarily during the spring and summer [95,121,123]. Chamise in mature chaparral is seldom browsed because stands are too dense for livestock or big game species to penetrate, making the current growth largely inaccessible [76,96,121,132]. Chamise sprouts on recently burned sites contribute greatly to total available forage within chaparral communities [9,76,113,121,123]. Sprouts provide a large volume of medium quality browse for domestic sheep and goats, and mule deer [9,34,48,121] but are browsed for only one to two postfire seasons because they rapidly become unpalatable [9,76,112]. Mule deer and domestic sheep frequently strip the leaves of sprouts, which are larger, more succulent, and less sclerophyllous than those of unburned plants [76,121]. Seedling leaves are eaten by domestic sheep and cattle [25]. Dusky-footed woodrats gather chamise leaves and bark and store them in dens for year-round consumption [55]. Chamise provides habitat for a variety of small and large wildlife species [96]. Dense stands serve as hiding, resting, and nesting sites for many smaller birds and mammals. Wirtz [132] compiled a list of common mammals, amphibians, reptiles, and birds associated with southern California chaparral. PALATABILITY : Chamise is largely unpalatable to most livestock and wildlife [121]; burning, however, greatly enhances its palatability [9]. Domestic goats often show a preference for chamise on recently burned chaparral [34,122,123]. Following goat depletion of California scrub oak (Quercus dumosa), the animals readily consumed chamise, preferring it over Eastwood manzanita (Arctostaphylos glandulosa) and desert ceanothus (Ceanothus greggii) [34,96]. Chamise browse has been rated useless for horses, useless to poor for cattle, and fair to good for sheep, goats, and deer [121]. Goats preferentially select the flower clusters [34]. NUTRITIONAL VALUE : The nutritive content of chamise is not particularly high. Chamise is a satisfactory source of digestible energy, but it is not high in digestible protein [121]. Crude protein values are highest in newly initiated leaves [121]. Seasonal trends in the crude protein content of the current growth (stems and leaves) are as follows [8]: Average % (oven-dry basis) winter (Dec.- Feb.) 7.0 spring (Mar.- May) 13.3 summer (Jun.- Aug.) 8.3 fall (Sept.- Nov.) 5.8 Sprouts contain nearly twice as much water, minerals, and crude protein per unit of total dry weight as does the current growth of mature plants [112]. Chemical composition (percent of oven-dry weight) of samples taken in August in Lake County is compared below [112]. young mature stems/ crown sprouts seeds formed silica-free ash 6.16 1.79 calcium 1.26 0.38 phosphorus 0.55 0.08 calcium:phosphorus 2.3 4.7 crude protein 17.53 2.89 crude fiber 12.89 28.71 moisture content 130 39 COVER VALUE : Chamise provides escape, bedding, resting, and thermal cover for mule deer [18,96]. In southern California chaparral, bighorn sheep prefer relatively open habitats on steep, south-facing slopes where the vegetation consists of a 30 percent cover of chamise, birchleaf mountain-mahogany (Cercocarpus betuloides), and chaparral whitethorn [12]. VALUE FOR REHABILITATION OF DISTURBED SITES : Chamise is suitable for revegetation because of its well-developed root system and drought resistance. Horton [52] listed it as useful for roadside erosion control plantings within chaparral. Recommended locations include sunny sites with deep or shallow soils at elevations between 500 and 3,500 feet (152-1,067 m) and sites with deep soils at elevations between 3,500 and 6,000 feet (1,067-1,829 m). Planting of pot-grown, bareroot stock produces satisfactory results [52]. Bareroot stock showed a 58 percent survival at the end of 2 years when planted at 2,700-foot (823 m) elevation in soils 2 to 6 feet (.6-1.8 m) deep; surviving plants reached heights of 4 to 6 feet (1.2-1.8 m) within 8 years. Two-year-old seedlings transplanted from a burn, however, had only 5 percent survival [52]. Plants may be propagated in flats from seed sown in winter or spring [52]. Seeds should be soaked in 10 percent sulfuric acid for 15 minutes prior to sowing. Within 4 to 6 months, seedlings usually reach heights of 2 inches (5 cm) and can be transplanted to pots. Most seedlings are ready for field planting after approximately 1 year in the pot stage [52]. Plants may also be propagated from green wood cuttings taken in the spring [128]. OTHER USES AND VALUES : Chamise was used by Native Americans for a variety of medicinal purposes. Chamise oils were used to treat skin infections, and an infusion of the bark and leaves was used for syphilis [18]. A binding agent for arrows and baskets was made from scale insects found on chamise plants [18]. OTHER MANAGEMENT CONSIDERATIONS : Browse tolerance: Access to new growth is greatly improved following fire, and chamise is fairly tolerant of concentrated use at this time [9,76]. Moderate cropping by deer prolongs the period of enhanced palatability by keeping sprouts in younger growth stages and by stimulating additional browse production [6]. Without browsing sprout bases become woody and are no longer preferred [6,9]. Continued close browsing, however, kills most plants within 2 to 3 years [9,10]. Chamise is heavily browsed on Santa Cruz and Santa Catalina islands. Decades of severe overgrazing by feral animals (pigs, sheep, and goats) has removed more palatable species and given chaparral stands an open, arborescent structure. Chamise exhibits a noticeable browse line and a trend towards increased trunk diameter, canopy coverage, and height [15]. Chamise produces few basal sprouts under such intense browsing and is very susceptible to eradication [15,87]. Herbicides: Chamise is sensitive to 2,4-D [14,37,63,97]. Plants exhibit a wide range of response to ammonium sulfide or benzoic acid application [37].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Adenostoma fasciculatum
GENERAL BOTANICAL CHARACTERISTICS : Chamise is a diffusely branched, resinous, native shrub [25,91,121] from 2 and 12 feet (0.6-3.5 m) tall [91]. Plants are unarmed, spreading, and branch very close to the ground [91,128]. The many slender stems are erect and generally lack permanent branches [38]. Young stems have reddish bark; bark becomes gray and shreddy with age [38,91]. Linear, needlelike leaves occur in alternate fascicles along the stem [18,121]. Leaves are 0.25 inch (0.6 cm) long, sharp-pointed, heavily sclerified, and evergreen [18,38,65]. The inconspicuous, bisexual flowers are white and occur in showy, 1- to 4-inch-long (2.5-10 cm) terminal clusters [22,121]. The fruit is an achene [91,128]. Although rooting habit is variable [79,84], roots are usually deeply penetrating, much branched, and widespreading [38,49]. The root system is extensive in relation to the crown [78,79]. Chamise typically develops several taproots which penetrate fractured rock to depths of 10 to 12 feet (3.0-3.7 m) [38]; extensive laterals originate from the lignotuber [49]. Longevity of chamise is estimated at 100 to 200 years [52,66,116]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Chamise reproduces sexually and vegetatively. Since chamise seed germinates at high rates only after fire, seedling recruitment and population expansion are fire dependent [68,69]. Canopy rejuvenation through the production of new basal sprouts occurs with or without the influence of fire [68,70]. Reproduction by seed: Onset of seed production occurs early in chamise, often by 3 years of age [27]. Seed production does not appear to decrease with age. Ninety-year-old shrubs generally produce substantially greater quantities of seed than those 20 years of age [66]. Seeds are dispersed during the summer [69]. Because the small achenes are not highly specialized for wind dispersal, most seeds fall near the parent plant [69]. Although the seed crop is abundant, the majority of seeds are not filled and viability is quite low, in some cases 0 to 4 percent [38,69,88,126]. Maximum seed production occurs following winters with above-average rainfall [3,38]. Chamise produces a dimorphic seed population composed of dormant as well as readily germinable seeds [16,126]. Dormancy is imposed by a more or less impermeable seedcoat. Heat from fire scarifies the seedcoat and stimulates germination [16,69,126]. Christensen and Muller [16] found that germination was enhanced when seeds were exposed to temperatures of 160 to 180 degrees Fahrenheit (71-82 deg C) for 15 minutes. Keeley [69] suggested that heat shock from fire and the presence of charate (charred wood) may act synergistically to stimulate germination. In laboratory studies, Keeley found that addition of charate significantly increased germination (11%) relative to controls (4%). Maximum germination (18%), however, occurred when heat-treated seeds were incubated in the presence of charred wood [69]. Black sage (Salvia mellifera) apparently inhibits chamise germination [131]. Under natural conditions, dormant seeds accumulate in the soil until stimulated by fire to germinate [66,126]. Chamise seeds are unpalatable and seedbanks apparently are not subject to heavy predation [111]. Consequently, chamise seed densities increase over time [133]. Seed density in the seedbank beneath 9-year-old stands was estimated at 2,000 seeds per square meter while in 85-year-old stands, seed density was approximately 21,000 seeds per square meter [132]. Abundant germination from soil-stored seed occurs during the first rainy season after fire; germination during the second year is uncommon [54,67,111]. Although emergent seedling populations are quite high [45], mortality is substantial during the first several years [39,54,120]. On sites in southern California, approximately 90 percent of the seedlings that germinated during March and April died within the first year [80]. Drought stress during late spring and summer is a major cause of first-year seedling mortality [59,86]. By the end of the second growing season, drought-induced mortality decreases as seedlings develop sufficient root biomass [87]. Taproots of newly germinated seedlings are barely 2 inches (5 cm) long by July, whereas taproots of 2-year-old seedlings range between 8 and 12 inches (20-30 cm) [80]. Small mammal herbivory contributes significantly to mortality [17], particularly in the fall [85]. First-year mortality due to rabbits may be as high as 25 percent [85]. Failure to establish may also be due to lack of suitable microsites and competitive interference [43,86]. On southern California burns, survival of first-year seedlings was not affected by the presence of residual shrubs or herbaceous perennials; annuals, however, significantly reduced seedling growth [80]. Many chamise plants die during subsequent years [48,120], but some survive [48,54]. Twenty-five years after a fire in central California, chamise resulting from seed were still growing and had reached an average height of 31.9 inches (80 cm) [54]. A portion of chamise seed germinates without fire scarification under favorable moisture and temperature conditions [126,133]. A study of the seedbank beneath an 85-year-old stand of chamise indicated that 20 percent of the chamise seedbank (density averaged 9,500 chamise seeds/sq m) was readily germinable [133]. Although initial establishment sometimes occurs without the influence of fire [35,54,101], seedling survival beyond the first year is extremely low and usually limited to areas recovering from human disturbance or overgrazing [135]. In mature chaparral, seedlings occasionally establish in canopy gaps, but successful establishment almost never occurs directly beneath the canopy [17,38,69,70,72,134]. Vegetative regeneration: Chamise rejuvenates its crown by continually producing new sprouts from an established lignotuber [48,69,70]. Following disturbances such as fire or cutting, chamise sprouts vigorously from surviving adventitious buds on the lignotuber [57,120]. SITE CHARACTERISTICS : Chamise is the most common chaparral species throughout the foothills and coastal mountains of California [13,38,39]. It is present in approximately 70 percent of California chaparral [13,39]. It is most often associated with hot, xeric sites [43] over a wide range of elevations, soils, latitudes, and distances from the coast [44]. In southern California it is a ubiquitous dominant on outwash plains, mesas, ridges, and dry, south- and west-facing slopes at elevations up to 6,000 feet (1,800 m) [18,35,38,52,91,100,121]. Sites supporting chamise commonly receive between 10 and 40 inches (250 and 1,000 mm) of annual precipitation, and have a temperature range from 32 to 100 degrees Fahrenheit (0-38 deg C) [48]. In the southern Coast Ranges, where average annual rainfall ranges from 16 and 20 inches (400-500 mm), chamise occurs abundantly on all slopes and exposures and grows on deep, fertile soils as well as shallow, rocky ones [48,121]. As precipitation increases farther northward, chamise is largely restricted to the poorer soils and the drier, more exposed sites [48,120]. Chamise occurs in both pure and mixed stands [38,39,120]. Nearly pure (>80%) stands of chamise are impenetrable and are referred to as "chamisal" [20,25,43]. Such stands usually have shallow, rocky soils with a southern aspect [35,53]. SUCCESSIONAL STATUS : Chamise is a long-lived, shade-intolerant shrub [66] which dominates lower elevation chaparral throughout much of California [20,42]. Disagreement exists over whether its dominance is a reflection of a climatic climax [5,20] or is a fire-induced subclimax [42]. Hanes [43,44] stated that chamise chaparral is unable to perpetuate itself in a vigorous condition without recurrent fire and terms it a true "fire-type vegetation". Chamise stands older than 60 years of age are sometimes termed " decadent" [39,48]. Old stands have low species diversity and produce little annual growth, with dead stem biomass far exceeding live stem biomass [40,43]. Stand stagnation has been attributed to the accummulation of biochemicals in the soil that inhibit decomposition, humification, and nitrification [40,94,131]. Limited nutrient availability, especially of nitrogen, may partially contribute to the decline of chamise chaparral [119]. Fire rejuvenates stagnant stands by removing phytotoxic substances from the soil, increasing the concentration of available nutrients, and stimulating sprouting of adults and germination of dormant seed [44]. Chamise is present soon after fire and remains present in all stages of succession. It achieves initial postfire dominance through vigorous sprout production and establishment of large numbers of seedlings [9,74,120]. Typical vegetal cover on 1-year-old chamise chaparral burns also includes a high percentage of herbaceous vegetation and the seedlings and sprouts of associated shrubs and subshrubs. As chamise seedlings and sprouts grow during the first postfire decade, herbaceous vegetation rapidly declines; likewise, subshrubs and short-lived shrubs are restricted to smaller and smaller openings [29,45]. A dense stand of chamise typically develops within approximately 8 to 10 years [42], with chamise frequently comprising one-third of total cover [39]. Stands often exhibit complete canopy closure by 22 years of age [116]. In pure stands of chamise in southern California, chamise may reach 25 percent, 50 percent, and 55 percent cover within 10, 40, and 70 years of fire, respectively [53]. Short-lived shrubs and herbaceous cover are largely lacking from undisturbed stands of chamise chaparral [116]. Chamise probably produces allelopathic toxins which inhibit germination and growth of other species [16,17]. During summer drought, chamise leaves accumulate water-soluble phenolics as a result of normal metabolic activity; fog drip and rain transport the toxins into the soil [43,83]. Competition for light may also be a factor controlling seed germination beneath mature stands [67,69]. Although chamise has only a limited ability to colonize disturbed areas [135], it is capable of pioneering broken rock surfaces and alluvial washes [43]. Chamise may invade woodlands where grass cover is sparse and sometimes invades productive soils following fire [48]. On sites with relatively deep soils, decadent chamise may be replaced by annual grasses [35]. SEASONAL DEVELOPMENT : Stem elongation occurs from February through May [2,130]. Shoot organization in chamise consists of short and long shoots and has been described by Jow and others [62]. New leaves appear in late January or February and continue to develop as shoots elongate [2]. New foliage is not limited to the current season's growth; short shoots remain active and produce leaves on 2- to 8-year-old branches [62]. Leaves are retained for two growing seasons [118]. Chamise produces nearly twice the amount of reproductive tissue as it does new stems and leaves [89]. In Sequoia National Park, flowers develop on the current year's growth in June followed by fruit development in July [2]. Fruit ripening and dispersal is completed by August. At this time, inflorescences die back and new growth becomes woody [2]. Although flower bud development and flowering occur at a time of decreasing water potential, reproductive growth is somewhat resistant to summer drought conditions. Water stored in the lignotuber allows chamise to maintain reproductive growth despite low water potentials [2]. Ample rainfall during the season directly preceding major growth activity increases the quantity of reproductive as well as vegetative growth [2,38]. Root growth: The period of root growth lasts considerably longer than the seasonal flush of shoot growth [78]. Fine roots may grow for 5 to 7 months [78]. Carbohydrate reserves: Onset of shoot growth is preceded by carbohydrate mobilization to the shoot apex and correlated with a decrease in the starch concentration of the roots and lignotuber [108]. Demand for nutrients during canopy and reproductive growth is quite high and by the end of the spring growth season, carbohydrate reserves in the roots and lignotuber are largely depleted [61]. During the summer, water stress-induced suppression of photosynthesis results in a reduction in carbohydrate availability at the shoot apex, and shoot growth ceases [1,5,38,108]. Cessation of growth is followed by a gradual increase in root starch reserves over fall and winter [61,81].

FIRE ECOLOGY

SPECIES: Adenostoma fasciculatum
FIRE ECOLOGY OR ADAPTATIONS : Following fire, chamise sprouts from dormant buds on the lignotuber [60,68,73]. The lignotuber has a stored supply of carbohydrates, nutrients, and water which support vigorous growth [60]. Chamise also produces abundant seedlings from fire-activated, soil-stored seed [16,68]. Chamise rapidly reoccupies the postfire community. Chamise possesses a number of adaptations that enhance its flammability [39,93,104,116]. These adaptations result in intense, fast-spreading, potentially large fires which have an increased probability of occurring as a stand matures [104]. Chamise chaparral produces fuel loadings capable of supporting a moderately intense fire within approximately 15 years [103]. Adaptations which enhance flammability are discussed below. Chemical: The chemical composition of foliage includes high energy ether extractives (waxes, resins, oils, terpenes, and fats) and inorganic minerals that affect pyrolysis of carbohydrates [111]. Ether extractives in the foliage increase burning rate because of their high heat content and may account for as much as 34 percent of the available heat content of chamise [104]. In older plants, a significant increase in the ether extractive content of 1- and 2-year-old leaf and stem tissues apparently contributes to the increased flammability of older stands [116]. Volatile, high energy essential oils on the leaf surface also ignite at low temperatures [115,116]. Physical: Structural characteristics produce rapid rates of energy release [21,116]. Approximately 60 percent of chamise stems are less than 0.5 inches (1.27 cm) in diameter [21]. Large amounts of small-stemmed material, distributed continuously from ground level throughout the multistemmed canopy, lend spatial continuity to the fuelbed and facilitate heat transfer [104]. Chamise also retains dead material in the crown [116]. As a stand ages, this material accumulates and within 30 years may account for 50 percent of the fuel loading [111]. Besides igniting easily and burning fast, dead fuels preheat live fuels, further increasing stand flammability [21,111]. Physiological: Chamise is most flammable in the fall [111]. Fuel moisture drops significantly during hot, dry weather and increases the concentration of extractive chemicals [115]. POSTFIRE REGENERATION STRATEGY : Tall shrub, adventitious-bud root crown Small shrub, adventitious-bud root crown Ground residual colonizer (on-site, initial community)

FIRE EFFECTS

SPECIES: Adenostoma fasciculatum
IMMEDIATE FIRE EFFECT ON PLANT : Chamise is a fire-sensitive species [136], and mortality may be substantial following fire [54,74,105]. Perennating buds are located just beneath the soil surface and are quite susceptible to fire damage [3,136]. Mortality patterns are related to season of fire, fire intensity and severity, and fire frequency [125]. The following photographs show top-killed chamise immediately after the 2003 Otay Mountain Wildfire in San Diego County and chamise sprouting in postfire year 2. Photos are courtesy of the San Diego Wildfires Education Project.
Burned chamise in Otay Mountain Wilderness Area, 2003.
Sprouting chamise in Otay Mountain Wilderness Area, 2005.
Close-up of chamise sprouting from the root crown in 2005.

Season of burning:  Season of fire affects chamise lignotuber survival.
Spring or summer fires may kill up to 50 percent of plants, while fall
fires result in relatively little mortality [3,74,124].  Differential
mortality is related to seasonal fluctuations in the carbohydrate
reserves of lignotuber and large roots.  High starch concentrations are
apparently necessary for the onset of sprouting [89,119].  Starch
concentrations may be insufficient to ensure sprouting when chamise is
burned in the late spring or summer, since carbohydrate reserves have
been depleted during spring growth [61,81].  Over the summer months,
however, starch reserves are recharged as carbohydrates are translocated
to the lignotuber, and most plants sprout following fall fires.  In dry
years, major carbohydrate mobilization does not take place; under these
conditions, spring or summer fires might produce lower level mortality
than in more "normal" years [116].

Fire intensity:  Mortality increases with increasing fire intensity.
Following low-, moderate-, and high-intensity June fires in old-growth
chamise in Sequoia National Park, approximately 46, 64, and 80 percent
of chamise plants died, respectively.  Seasonal patterns of fire
mortality are further accentuated by differences in fire severity
associated with spring/summer versus fall fires.  Early season fires
move slowly through a stand and the downward heat pulse is greater than
that produced by rapidly carrying, fall fires.  As a result, fall fires
are generally less severe than spring/early summer fires and produce
less mortality.  In one case, a moderately intense spring (June) fire
resulted in 64 percent chamise mortality, whereas a moderately intense
fall (October) fire resulted in only 14 percent mortality [116].

Fire frequency:  Chamise is extremely susceptible to short-interval
fires.  High mortality of both seedlings and sprouts is likely when
fires recur on burns seeded to annual grasses.  Chamise density
(seedlings and sprouted individuals) was reduced up to 97 percent
following a grass fire on a 1-year-old burn [136].  Chamise seedlings
are more sensitive to frequent fire than sprouted plants [136].  In
northern California, Hedrick [48] reburned 1-, 2-, and 3-year-old burn
sites which had been seeded to grass and mustard.  Mortality of
established chamise was 77 percent on the 1-year-old burns, 24 percent
on the 2-year-old burns, and 34 percent on the 3-year-old burns.
Seedling mortality was 99, 98, and 100 percent on the 1-, 2-, and
3-year-old burns, respectively.  Fires occurring at very short intervals
may completely eradicate postfire seedling reproduction if the soil seed
reserve is not well established and reproductive maturity has not been
reached [58,136].  Although sprouts are generally capable of heavy seed
production by the second year after fire [65], chamise seeds exhibit
poor viability.  A number of years are required to build up the
seedbank.

Seedbanks:  Chamise seed is sensitive to high temperatures [16,40,130].
Depending on fire intensity and seed position in the soil, a large
portion of the seedbank may be destroyed as the soil temperature rises
during burning [111].  While abundant seed is present beneath the shrub
canopy and in gaps between shrubs, burning modifies the seedbank by
concentrating readily germinable seed in the shrub interspaces.  Soil
temperatures during burning are lower in the shrub interspaces, and more
seeds survive fire in these interspaces than below the canopy [23].
Fire sensitivity is increased if seeds have imbibed water [23,99], and
seed mortality is high following spring fires, which are often severe
[111].  Reduced seed mortalities can be expected when fires occur under
dry soil conditions associated with late summer and fall.


DISCUSSION AND QUALIFICATION OF FIRE EFFECT : 
Small plants with a prefire biomass between 2 and 11 pounds (1 and 5 kg)
are particularly prone to fire mortality [3] because of their shallow
lignotubers and presumably smaller carbohydrate reserves [3,116].
Individuals with larger lignotubers are generally more fire tolerant,
although large shrubs which have survived previous fires may have more
dead material in their crowns, making them more prone to fire mortality
[124].  Fire susceptibility of larger plants also increases in older
stands where high fuel loads produce severe fires [74].  Mean lignotuber
area of fire-killed plants in a 90-year-old stand was 35 square inches
(227 sq cm) compared to 12 square inches (79 sq cm) for plants in a
23-year-old stand [74].  Very large chamise lignotubers tend to rot in
the center and are less capable of sprouting after fire [60].  While
young seedlings are readily killed by most fires, fire tolerance
increases with age.  In southern California, large numbers of 4-year-old
chamise sprouted following intense summer grassfires [58,136].


PLANT RESPONSE TO FIRE : 
Postfire regeneration in chamise involves a combination of sprout
regeneration and seedling recruitment [48,54,69,120].

Vegetative regeneration:  Although considerable plant mortality may
occur following fire, at least some percentage of the chamise population
survives and sprouts [39,42,61,120].  Residual plants typically sprout
within 6 weeks of fire regardless of season [108].  On sites in southern
California, chamise along moist ravines sprouted within 10 days of a hot
July wildfire [105].  Sprouts originate on the lignotuber from a narrow
band of tissue located 0.2 to 1 inch (0.5-2.5 cm) below the soil surface
[109].  Perennating buds along the periphery of the lignotuber are the
first to initiate sprouts, followed by buds near the base of charred
stems [6].  Sprout production varies by lignotuber size.  Large plants
usually possess large lignotubers which produce as many as 500 sprouts
[6].  During the first year sprout numbers are drastically reduced as
larger stems gain dominance [6,111].

Sprouts use stored carbohydrate reserves to achieve rapid growth
[109,111].  Rapid shoot elongation typically occurs during the first
spring following fire [60,108].  After fires in northern California,
sprouts averaged 20 inches (50 cm) by the end of the first postfire
growing season [9].  Shrubs with a large prefire biomass typically
produce the most vigorous sprouts and can be expected to dominate the
postfire community [124].  Baker and others [3] indicated that sprout
biomass of residual plants at the end of postfire year 1 is positively
correlated with prefire biomass.  Stem growth slows during postfire
years 2 and 3 [54,108], and growth during subsequent years declines
until it is almost negligible by 20 years after fire [54,108].  Horton
and Kraebel [54] reported that 5-year-old sprouts reached an average
height of 33.6 inches (84 cm), while 20-year-old sprouts were only 40.4
inches (101 cm) tall [54].

The pattern of postfire sprout growth usually follows that of mature
plants.  Time of fire, however, may alter the initial pattern of
postfire shoot growth.  On sites in the southern Sierra Nevada, plants
burned in late June or early August produced sprouts that grew
continuously until the second postfire summer [108].  The reduced leaf
area of sprouted plants limited transpiration losses and resulted in
higher shoot water potentials, permitting shoot growth through the
summer drought period [46,47].  While summer fires (at a time of reduced
carbohydrate reserves) initially resulted in significantly shorter
plants, shoot heights of plants burned in different seasons were similar
by the end of the second postfire year [108].

Seedling regeneration:  Chamise produces an abundant crop of seedlings
from soil-stored seed [17,39,45].  While a flush of initial seedling
establishment may occur immediately following fire, subsequent mortality
is quite high [59,73,85].  On sites in southern California, seedling
densities in March ranged from 91,427 to 180,383 seedlings per acre
(37,000-73,000 seedlings/ha) but dropped to 29,652 to 34,594 pere acre
(12,000-14,000/ha) by June [73].  The degree to which seedlings
contribute to the postfire recovery of chamise is quite variable and
appears related to site conditions, amount of fire-induced adult
mortality, and stand age [39,58].  On sites where the majority of plants
survive fire, prefire shrub density is maintained and little seedling
establishment occurs.  Conversely, seedling establishment is often
substantial and critically important in regaining prefire levels on
sites where adult survival is low [3,66].

Recovery:  Because of hot, dry site conditions, postfire growth of
chamise chaparral is slow compared to other chaparral types [53,100].
Initial sprouting response may be substantially reduced following
intense summer fires, since more of the meristematic tissue in the
lignotuber is killed [116].  Four months after a July wildfire in
southern California, chamise plants produced up to 12 sprouts per plant,
but sprouts rarely exceeded 12 inches (30 cm) in length [105].  In
southern California chamise communities, chamise rapidly dominates the
postfire community and commonly comprises at least 33 percent of the
vegetation on 10-year-old burns.  In stands 22 to 40 years of age, it
reaches a maximum of approximately 50 percent of the total vegetative
cover [39].


DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : 
Since chamise sprouts following fire, reproduction from seed is
generally considered facultative [43,68].  Chamise relies primarily on
vegetative reproduction for postfire establishment on foothill sites in
the southern Sierra Nevada [116,118].  Although as many as 2.73 million
seedlings per hectare may emerge following fire, they are usually
outcompeted by faster growing sprouts [117].  Likewise, vegetative
reproduction is the predominant mode of postfire regeneration at higher
elevations in southern California mountains [71,73,129].

As chaparral sites become increasingly arid, however, sprouting tends to
be less successful and seedling recruitment more prevalent following
fire [39,136].  On droughty, low elevation sites in southern California,
chamise depends to a large degree on successful seedling establishment
for population replacement after fire [56,58,71,72,136].  Howe and
Carothers [58] found that chamise seedlings grew vigorously and
contributed significantly to postfire stands at elevations between 1,312
and 1,968 feet (400-600 m) near Newhall, California, in Los Angeles
County.  Chamise seedlings comprised approximately 86 percent of the
chamise population on 6- to 9-year-old-burn sites.  Although seedlings
grew more slowly than sprouts during the first few postfire seasons,
they reached heights equal to that of sprouted plants within 8 to 9
years.  On 6-year-old, north-facing burns, however, vegetative
reproduction was the predominant mode of regeneration [58].  Hanes [39]
indicated that altitude also influences mode of postfire reproduction.
He found that seedlings comprised a higher proportion of the postfire
vegetation on burn sites between 1,000 and 2,000 feet (300-600 m) than
between 2,000 and 4,000 feet (600-1,200 m).


FIRE MANAGEMENT CONSIDERATIONS : 
Fire frequency:  Chamise is adapted to a fire cycle range from 10 to 100
years.  It can regenerate after fire intervals of over 100 years,
however [68,74,90,116].  Its capacity for canopy rejuvenation without
fire allows chamise to persist through long fire-free intervals.
Stohlgren and Rundel [125] suggested 30 to 80 years as a "typical" fire
frequency for chamise chaparral communities in Sequoia National Park.

Influence of ryegrass seedings:  Ryegrass (Lolium multiflorum) is often
seeded onto recently burned chaparral as a means of emergency
revegetation [4].  Ryegrass, however, inhibits growth and development of
chamise seedlings [33], and ryegrass substantially reduces postfire
chamise seedling establishment [4,33].  On seeded burns in southern
California, almost no chamise seedlings established where first-year
ryegrass cover ranged between 40 and 90 percent [19].

Ryegrass seedings also produce an easily ignitable fuel bed that
increases the likelihood of an early reburn.  Fires occurring at short
intervals have the potential to cause significant changes in species
density and composition within chamise chaparral [4].  Not only do
frequent fires produce high mortality of sprouted plants [136], but
postfire seedlings (derived from the previously dormant seedbank) are
also killed, thereby depleting the on-site seed reserve [66,136].
Consequently, chamise is unable to reestablish, and gaps in the shrub
matrix are subject to invasion by coastal sage scrub species such as
black sage, California sagebrush, and California buckwheat.  The site
may be dominated by coastal sage scrub species for 100 years or more
[4].

Deer browse:  Deer use of chamise is often extensive immediately
following fire [9,113,121].  Browse value of sprouts lasts for only 2 to
3 years because plants quickly mature to less nutritious stages or die
from overuse [96].  To enhance deer use of sprouts, cattle access to
burns should be restricted during the first postfire season [113].
Because of the lack of adequate escape cover, only the periphery of large
burns receive extensive deer use prior to the second postfire season.
The center of large burns are rarely if ever utilized during the first
several seasons [96].  Close utilization within the first year may kill
chamise, and mortalities of up to 64 percent are possible under intense
browsing pressure [10].

Late winter or early spring fires are most favorable for production of
deer browse because succulent sprouts with a high nutrient value are
produced almost immediately, and subsequent sprout growth is rapid
during the spring growth period [76]. If fires are conducted after
mid-September in northern California, sprouting may not be profuse until
the following spring [9].  Fires resulting in total plant consumption
produce the most usable browse, since deer tend to avoid burned chamise
with main scaffold branches remaining [9,24].

REFERENCES

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