Bromus rubens, Bromus madritensis
AUTHORSHIP AND CITATION:
Simonin, Kevin A. 2001. Bromus rubens, Bromus madritensis.
Simonin, Kevin A. 2001. Bromus rubens, Bromus madritensis.
In: U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Laboratory (Producer).
Available: http://www.fs.fed.us/database/feis/ .
Red brome –
Anisantha madritensis (L.) Nevski 
B. madritensis ssp. rubens (L.) Husnot [41,46]
NRCS PLANT CODE :
There is taxonomic disagreement about foxtail chess (Bromus madritensis L.) (Poaceae) in the
strict sense. Following the systematics of several authorities, this report uses the scientific name
Bromus madritensis L. for foxtail chess and Bromus rubens L. for red brome [36,47,49,80,82,83].
Some systematists recognize these entities as subspecies of foxtail chess:
the type subspecies (Bromus madritensis ssp. madritensis) and
red brome (Bromus madritensis ssp. rubens (L.) Husnot) [41,46].
There is consensus that the 2 entities are very closely related [1,84], with both
scientific names used in current literature.
This report considers Bromus madritensis in the broad sense. Studies concerning United States populations
of B. rubens may be interpreted as B. m. ssp. rubens [1,84].
Where possible, distinctions are made between foxtail chess (B. madritensis)
and red brome (B. rubens). The common name foxtail chess is used
when discussing the species as a whole, and when literature cited does not distinguish between
foxtail chess and red brome in areas where their distributions overlap (mostly California).
Red brome refers to B. rubens.
FEDERAL LEGAL STATUS:
No special status
The California Exotic Pest Plant Council includes foxtail chess on the A-1
list: Most invasive wildland pest plants; widespread .
DISTRIBUTION AND OCCURRENCE
SPECIES: Bromus rubens, Bromus madritensis
Foxtail chess is native throughout Europe and the British Isles
[36,41,46,49,76,80,83]. In western North America, it occurs widely in coastal
and central California and is also recorded from Reno,
Nevada, and Portland, Oregon [66,84]. It has a scattered distribution to the
east, occurring in Michigan, Virginia, and Mississippi . It is also
introduced in Hawaii .
Red brome is native to southern Europe . In North America, it is
distributed from central Washington south to Baja California and east to central
Idaho, southwestern Texas, and Sonora [66,84]. It is casually adventive in the
Northeast  and introduced in Hawaii .
provides distributional maps of foxtail chess and of red brome.
FRES28 Western hardwoods
FRES30 Desert shrub
FRES34 Chaparral-mountain shrub
FRES40 Desert grasslands
FRES42 Annual grasslands
STATES/PROVINCES: (key to state/province abbreviations)
BLM PHYSIOGRAPHIC REGIONS :
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
KUCHLER  PLANT ASSOCIATIONS:
K027 Mesquite bosques
K030 California oakwoods
K035 Coastal sagebrush
K038 Great Basin sagebrush
K043 Paloverde-cactus shrub
SAF COVER TYPES :
249 Canyon live oak
250 Blue oak-foothills pine
255 California coast live oak
SRM (RANGELAND) COVER TYPES :
201 Blue oak woodland
202 Coast live oak woodland
205 Coastal sage shrub
206 Chamise chaparral
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
209 Montane shrubland
211 Creosotebush scrub
214 Coastal prairie
215 Valley grassland
401 Basin big sagebrush
413 Gambel oak
414 Salt desert shrub
503 Arizona chaparral
507 Palo verde-cactus
HABITAT TYPES AND PLANT COMMUNITIES:
Foxtail chess and red brome are invasive weeds. They are most prevalent in desert shrub and annual grassland communities
California: Foxtail chess is a dominant invader of the historical perennial grasslands of California
[2,5]. It is widely distributed over central and southern California . It has also invaded
steppe ecosystems of the southern San Joaquin Valley, the Great Basin,
and the Mojave Desert .
Common grass associates of foxtail chess within the western foothill annual
grassland of the Sierra Nevada
soft chess (Bromus mollis), ripgut brome (B. diandrus),
foxtail fescue (Vulpia myuros), slender wildoat (Avena barbata), California melic
(Melica californica), and Sandberg bluegrass (Poa secunda).
Common forbs include cutleaf filaree (Erodium cicutarium), Napa thistle (Centaurea melitensis),
clover (Trifolium spp.), hairy lotus (Lotus subpinatus), and godetia (Godetia dudleyana) .
Common associates in California oak savannah include blue oak (Quercus douglasii),
interior live oak (Q. wislizenii) [27,32], valley oak (Q. lobata) ,
gray pine (Pinus sabiniana) [27,32], and
California buckeye (Aesculus californica) . Common shrub associates include
wedgeleaf ceanothus (Ceanothus cuneatus) and chaparral whitethorn (C. leucodermis).
Soft chess, ripgut brome , slender wildoat,
wild oat (A. fatua) [32,78], mouse barley (Hordeum leporium) , and
fescue (Festuca spp.) 
are common grasses. Common forb
associates include cutleaf filaree [27,32], longbeak storksbill
(Erodium botrys), burclover (Medicago polymorpha), and clover .
Common coastal sage scrub shrub associates of foxtail chess include black sage (Salvia mellifera) and
California buckwheat (Eriogonum fasciculatum) .
Classifications describing plant communities in
which foxtail chess is a dominant species are as follows:
Nevada: Red brome is a dominant annual of blackbrush (Coleogyne
ramosissima) communities at 4,000 to 5,000 feet (1,200-1,500 m).
In the western Yucca Flat of southern Nevada, red brome is a member of creosotebush (Larrea tridentata)-blackbrush
transition communities between 4,000 and 4,300 feet (1,200-1,300 m), and is infrequent in creosotebush
associations between 3,000 and 4,000 feet (900-1,200 m). Red brome is abundant within
creosotebush-saltbush (Atriplex spp.)-blackbrush communities . Red brome is a prominent
member of blackbrush communities within the Mojave Desert [11,20].
Utah: Red brome is common in blackbrush, juniper (Juniperus spp.), creosotebush , and sand sagebrush (Artemisia filifolia) [64,70]
communities of Utah. It is most commonly found at base of shrubs in shadscale and white burrobush (Hymenoclea salsola)
Arizona: Red brome is an invader of Arizona desert scrub . It is also an invasive understory
species of mature mesquite bosques (Prosopis spp.),
which were historically open with a ground cover of saltbushes or annual and
perennial native grasses and forbs .
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Bromus rubens, Bromus madritensis
GENERAL BOTANICAL CHARACTERISTICS:
Foxtail chess and reed brome are a cool-season  exotic annual grasses with culms from 4 to 28 inches (10-70 cm) tall. Their inflorescence is a panicle, 1 to 4 inches (3-11 cm) tall,
with long awns . Foxtail chess is distinguished from red brome by having relatively hairier stems and leaf sheaths, more lax
panicles, and wider
Foxtail chess is drought resistant, with high water-use efficiency [10,21].
RAUNKIAER  LIFE FORM:
Foxtail chess germinates well under the winter temperature regime of southern California .
Sunlight may enhance germination at higher temperatures. Freshly harvested red brome
seeds are unable to germinate in the dark at temperatures
above 59 degrees Fahrenheit (15 °C), but freshly harvested seeds usually germinate in the dark
at temperatures between 41 and 59 degrees Fahrenheit (5-15 °C). White light inhibits germination
even at low irradiance. Dry storage at 68 degrees Fahrenheit (20 °C) removes the inhibitory
effects of higher temperatures ( > 68 degrees Fahrenheit (20 °C)), widening the
seeds' temperature range for germination and eliminating their sensitivity to light .
Red brome germinates at a less exact rainfall and temperature requirement than native species .
In general, fall rains promote germination and establishment .
In blackbrush communities of southern Nevada, Beatley  found red brome germination
followed heavy rains (> 1 inch (3 cm)) between October and December. However, germination is not limited
to fall and may occur following heavy spring rains.
Laude  evaluated seedling emergence from freshly harvested and stored red
brome seeds, which were were planted in a greenhouse setting and monitored for 28 days. Increased
storage time increased percent emergence, with germinants achieving 100% emergence in October,
5 months after mid-June seed collection.
High levels of cadmium and nickel impair emergence of red brome .
Foxtail chess and red brome prefer disturbed sites in Mediterranean climates [41,66]. Elevational ranges
are described below by state:
foxtail chess, red brome
|| 7,200 feet (2,200 m) 
|| 1,200 to 6,000 feet ( 370-1,800m) [44,48]
|| 4,000 to 5,000 feet (1,200-1,500 m) 
||3,000 to 5,000 feet (910-1,520 m) 
Regional: In California, foxtail chess prefers areas receiving less than 9.8 inches (250 mm)
annual rainfall . Foxtail chess is a dominant species in California valley grasslands receiving
less than 7.5 inches (190 mm) rainfall , and is abundant in California valley grasslands receiving
less than 12 inches (305 mm) annual precipitation. In areas
with annual precipitation greater than 12 inches (305 m), foxtail chess is replaced by soft chess (Bromus mollis)
Bowers  monitored the relative abundance of northern Mojave Desert annuals over 6 years
in relation to precipitation. Red brome density was highest during years
receiving 2.4 to 4.2 inches (63-109 mm) precipitation; results are summarized below:
Pre-census precipitation (mm) between 1970-1976:
||Mean red brome/0.25m2
Soils: Red brome commonly
occurs in small patches on shallow soils, growing best where there is little competition from other annuals .
In southern Nevada, red brome occupies blackbrush communities with coarse-textured
soils, showing best growth under shrubs and peripheries of shrub canopies . Upland clay and sandy loam
ranges and rolling sandy hills receiving
8 to 12 inches (203-305 mm) precipitation promote good growth in southern Utah
Red brome is often found in areas with relatively high levels of sulfur dioxide
The presence of foxtail chess and red brome are closely related to annual precipitation.
They are commonly early or mid-seral species where annual
precipitation is greater than 9.8 inches (250 mm) .
Red brome is commonly an early to mid-seral species in California chaparral. It is usually sparse in early succession
of northern California but may increase rapidly in areas of low soil fertility and moisture .
population numbers require several years for seed dispersal into burns or buildup from on-site producers.
Continued disturbance such as grazing and repeated low-severity fires favor red brome over native
early-seral chaparral species .
Within blackbrush communities of Nevada, red brome persists in high-density stands for many
In general, red brome initiation and establishment is a direct response to fall rains. Initial
growth is relatively slow, followed by a rapid increase in vegetative growth coinciding with warming spring
temperatures . Flowering and fruiting generally
occur in April and May . Seeds are disseminated in summer .
No profound phenological differences are apparent between red brome individuals from the Mediterranean
and California regions . However, Wu and Jain  have observed phenological variation in seed weight,
lemma length, plant height, and tiller production between populations
of different environments.
SPECIES: Bromus rubens, Bromus madritensis
FIRE ECOLOGY OR ADAPTATIONS:
Red brome generally shortens fire return intervals [59,60,87].
The increased presence of red brome
has promoted fires in areas where fire was previously infrequent due to insufficient
. Once established red brome may increase fire frequency by enhancing potential for start
and spread . In general, red brome produces an abundant and continuous cover of persistent fine fuels,
promoting fast, "hot" fires .
Desert: Areas of the Mojave Desert dominated by red brome are more susceptible to
fire than areas dominated by native forbs. Dead red brome culms and blades are persistent
(commonly 2 years); herbage of most Mojave Desert annual species usually lasts 1
year or less.
Red brome produces high amounts of persistent flammable fuels in perennial
plant interspaces, promoting ignition and spread .
Heat generated by burning red brome is sufficient to ignite and consume dead stems of native
Mojave Desert forbs. Flames may
also consume small shrubs such as white bursage (Ambrosia dumosa), winterfat (Krascheninnikovia lanata),
white burrobush, and Anderson wolfberry (Lycium andersonii).
However, flames fueled by red brome are generally insufficient to ignite large shrubs such as
creosote bush .
Within the Sonoran Desert, dead and dry red brome is easily ignited, supporting fast-moving surface fires . Fire return
intervals are also shortened, changing the vegetal composition through increase of non-native
components and loss of native plant species .
The invasion of red brome has contributed to short fire return intervals and the subsequent degradation of
chaparral . Keeley  has observed shorter fire intervals (< 10 years) resulting from the increased
presence of red brome and other exotic annuals. Greater fire frequency has promoted
the degradation of native herb communities and promoted communities of chaparral shrubs with an exotic,
annual understory . Dead stems and litter are persistent, promoting spread of
fire in shrub communities, especially across areas between shrubs and trees [44,87].
O'Leary and Westman  attribute red brome within early postfire coastal sage scrub
communities to off-site seed sources.
As nonnative species, foxtail chess and red brome have no historic fire regime in North America.
The following table provides some fire regime intervals for areas where foxtail chess
and red brome presently occur. Foxtail chess and red brome may alter fire intervals within these communities. Find further fire regime information for the plant communities in which these
species may occur by entering the species' names in the FEIS home page under "Find Fire Regimes".
||Adenostoma and/or Arctostaphylos spp.
||< 35 to < 100
||Artemisia tridentata/Pseudoroegneria spicata
||< 35 to < 100
|California montane chaparral
||Ceanothus and/or Arctostaphylos spp.
||Cercidium microphyllum/Opuntia spp.
||< 35 to < 100
|mountain-mahogany-Gambel oak scrub
||Cercocarpus ledifolius-Quercus gambelii
||< 35 to < 100
||< 35 to < 100
||< 35 to < 100
||Larrea tridentata-Leucophyllum frutescens-Prosopis glandulosa
|oak-juniper woodland (Southwest)
||< 35 to < 200
|blue oak-foothills pine
||Quercus douglasii-Pinus sabiana
|interior live oak
||< 35 
POSTFIRE REGENERATION STRATEGY :
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)
SPECIES: Bromus rubens, Bromus madritensis
IMMEDIATE FIRE EFFECT ON PLANT:
Fire kills foxtail chess and red brome .
DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
PLANT RESPONSE TO FIRE:
Red brome establishes from on- or off-site seed sources following fire .
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Red brome showed dramatic increases in the Sonoran Desert following prescribed burns
in a desert scrub of paloverde, buckhorn cholla (Opuntia acanthocarpa), and
Results are summarized below :
Cover (%) within open microhabitat:
||12 postfire months (1985)
||24 postfire months (1985)*
||36 postfire months (1986)*
Cover (%) in shrub microhabitat:
*Coverage data taken from areas burned in 1983
||12 postfire months (1985)
||24 postfire months (1985)*
||36 postfire months (1986)*
Davis and Hickson  evaluated postfire vegetation
development within coastal chaparral of California. Past fires in the area studied was
either accidental or
planned burns. Areas sampled ranged from 1 to 50 years since fire. Overall, red
brome showed greater frequency of occurrence within oak plots than in
Red brome was an important invader of chamise chaparral in the 3rd and 5th postfire seasons,
slowly disappearing with increased brush cover. Before fires, red brome was restricted to
trails, firebreaks, and openings .
Red brome was a dominant species on western and southern slopes (330 feet (100 m)) the first
2 years after fire in California coastal sage scrub. Postfire annual precipitation was 1 to 5 times
greater than mean annual precipitation .
Fall burns: Red brome and California buckwheat were dominant species 5 years
after a September wildfire in a 7-year-old rockrose (Cistus spp.) field, originally
planted from nursery grown seedlings. The wildfire occurred on a 30 to 35 degree north facing slope at
2,700 feet (820 m). .
Hansen  observed postfire coverage of red brome after fall prescribed burns in the Tulare
Basin of California.
Seasonal precipitation directly influenced red brome postfire response. One-year postfire
cover of red brome was equal to or greater than control plot coverage when precipitation was below average and
the majority of precipitation occurred in the fall (September-November). In contrast, red brome
presence was greatly reduced in the 1st and 2nd postfire growing season if greater than
normal precipitation occurred in fall and spring. Repeated burning had the greatest negative
effect on red brome numbers, resulting in large decreases even in wet years.
Spring burns: Cave  monitored 1- and 2-year postfire vegetation response in the
Sonoran Desert after a 12 June 1981 controlled burn and a 26 April 1980
wildfire. Sites were at 1,500 feet (450 m) on a paloverde (Cercidium microphyllum)-cactus (Opuntia
spp.)-triangle bursage association. Red brome density was reduced by 96 to 98%
1st postfire year compared to unburned and prefire controlled burn sites, respectively. Biomass
was reduced by 70 and 65%, with cover showing 86 and 80% reductions.
Precipitation prior to and during the 1982 growing season was less than
precipitation in 1981.
Red brome dominated blackbrush areas 1 year after a "hot" spring wildfire .
For information on response of red brome and other annual grassland herbs
to prescribed fire, see Hansen's  thesis:
The effect of fire and fire
frequency on grassland species composition in California's Tulare Basin.pdf.
FIRE MANAGEMENT CONSIDERATIONS:
O'Leary and Westman  suggest off-site seed as a source of red brome within early postfire coastal sage scrub
Stipa spp. have successfully competed with red brome within perennial grasslands of Camp Pendleton,
California, subjected to annual burning .
Shrublands: Within the Mojave Desert of Arizona, red brome prefers disturbed sites, especially
areas where shrubs have been removed by fire.
Red brome readily invades blackbrush communities susceptible to fire. Once established
red brome increases fire frequency by enhancing the potential for start and spread .
Red brome shows vigorous vegetative growth in blackbrush communities where shrubs have been removed by
fire. Red brome is prominent the first 2 to 3 postfire years in blackbrush communities,
after which perennial grasses and shrubs dominate .
Wildfires in California chaparral occur during hot dry summer months.
Prescribed burning usually occurs during winter months, resulting in low-intensity burns. Red brome is a dominant
species following winter fires. Temperatures of low-intensity winter fires are
not high enough to kill red brome seeds .
Fire spread is extensive and rapid when red brome is codominant with
Schismus spp. in native perennial
interspaces of the Mojave Desert. Data for experimental fires conducted in August of 1995 are presented below. Fine fuels are presented in
kg/ha, dominant annuals are those with > 25% relative cover .
For further information on this study, see the Research Project Summary Nonnative annual grass fuels and fire in
California's Mojave Desert.
||Beneath canopy fuel
||Dominant annuals, beneath canopy
||Dominant annuals, interspace
||Interspace fire spread (m/minute)
||Area burned (% of 2.25 ha)
SPECIES: Bromus rubens, Bromus madritensis
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
In general, foxtail chess and red brome are of little value to livestock and big game. The long awns of red brome
are harmful to livestock when seeds are ripe . Under some circumstances, red brome
may provide a source of forage. Krausman and others  observed light desert mule deer
use (< 1% of seasonal diet)
of red brome in Arizona. Desert bighorn sheep also use red brome .
Desert cottontail prefer red brome. Heaviest use occurs in winter .
The palatability of red brome to livestock and
wildlife in Utah has been rated as follows :
|Upland game bird
|Small nongame bird
The calcium:phosphorous ratio for red brome is 1:2 .
Seegmiller and others  conducted a nutritional analysis of red brome within the Harlequin Mountains, Arizona. Red
brome parts selected for analysis (leaves, flowers, new growth) were those used by desert bighorn sheep. Results (mean %)
are presented below:
Foxtail chess provides fair cover for small mammals and small nongame birds .
VALUE FOR REHABILITATION OF DISTURBED SITES:
OTHER USES AND VALUES:
OTHER MANAGEMENT CONSIDERATIONS:
Red brome is common in open canopies of Arizona chaparral subjected to heavy grazing .
Betancout  attributes red brome expansion in the Upper
Sonoran Desert of central and southern Arizona to climate change. Since 1976, increased winter precipitation has promoted the
spread of red brome.
In relatively dry areas of the Southwest, red brome may displace native species during wetter years [4,14,44].
Relatively drier winters and wetter summers may slow the red brome invasion .
The smut Ustilago bullata is common on red brome in Nevada. Infected plants produce
fewer viable seeds than uninfected plants .
In California chaparral, fall germination of foxtail chess gives the grass a competitive
advantage over shrub seedlings, which usually germinate in spring .
Foxtail chess competes with California sagebrush (Artemisia californica)
seedlings during their 1st growing season. Planting shrubs that are 1 or more
years old increases their competitive ability against red brome . Bartolome and others  found mulch
provided no significant (p > 0.05) benefit to standing crops foxtail
areas of California receiving less than 9.8 inches (250 mm) annual precipitation.
Foxtail chess is an annual weed in California cereal crops .
Bromus rubens, Bromus madritensis: REFERENCES
1. Allred, Kelly. 2001. [E-mail to Janet Howard]. January 3. Las Cruces, NM: New Mexico State University. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab, Missoula, MT; RWU 4403 files. 
2. Amme, David; Pitschel, Barbara M. 1990. Restoration and management of California's grassland habitats. In: Hughes, H. Glenn; Bonnicksen, Thomas M., eds. Restoration `89: the new management challenge: Proceedings, 1st annual meeting of the Society for Ecological Restoration; 1989 January 16-20; Oakland, CA. Madison, WI: The University of Wisconsin Arboretum, Society for Ecological Restoration: 532-542. 
3. Ashby, William C.; Hellmers, Henry. 1955. Temperature requirements for germination in relation to wild-land seeding. Journal of Range Management. 8: 80-83. 
4. Banner, Roger E. 1992. Vegetation types of Utah. Journal of Range Management. 14(2): 109-114. 
5. Barry, W. James. 1972. The Central Valley prairie. Vol.1. Sacramento, CA: State of California, Department of Parks and Recreation. 82 p. 
6. Barry, W. James. 1985. Ecosystem restoration in the California state park system. In: Rieger, John P.; Steele, Bobbie A., eds. Proceedings of the native plant revegetation symposium; 1984 November 15; San Diego, CA. San Diego, CA: California Native Plant Society: 22-33. 
7. Bartolome, James W. 1987. California annual grassland and oak savannah. Rangelands. 9(3): 122-125. 
8. Bartolome, James W.; Stroud, Michael C.; Heady, Harold F. 1980. Influence of natural mulch on forage production on differing California annual range sites. Journal of Range Management. 33(1): 4-8. 
9. Bates, Patricia A. 1983. Prescribed burning blackbrush for deer habitat improvement. Cal-Neva Wildlife Transactions. [Volume unknown]: 174-182. 
10. Beatley, Janice C. 1966. Ecological status of introduced brome grasses (Bromus spp.) in desert vegetation of southern Nevada. Ecology. 47(4): 548-554. 
11. Beatley, Janice C. 1974. Phenological events and their environmental triggers in Mojave Desert ecosystems. Ecology. 55: 856-863. 
12. 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. 
13. Betancourt, Julio L. 1996. Long- and short-term climate influences on Southwestern shrublands. In: Barrow, Jerry R.; McArthur, E. Durant; Sosebee, Ronald E.; Tausch, Robin J., compilers. Proceedings: shrubland ecosystem dynamics in a changing environment; 1995 May 23-25; Las Cruces, NM. Gen. Tech. Rep. INT-GTR-338. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 5-9. 
14. Billings, W. E. 1949. The shadscale vegetation zone of Nevada and eastern California in relation to climate and soils. The American Midland Naturalist. 42(1): 87-109. 
15. Biswell, H. H.; Gilman, J. H. 1961. Brush management in relation to fire and other environmental factors on the Tehama deer winter range. California Fish and Game. 47(4): 357-389. 
16. Biswell, Harold H. 1974. Effects of fire on chaparral. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and ecosystems. New York: Academic Press: 321-364. 
17. Bjorndalen, Jorn Erik. 1978. The chaparral vegetation of Santa Cruz Island, California. Norwegian Journal of Botany. 25: 255-269. 
18. Bolander, Donald H. 1982. Chaparral in Arizona. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 60-63. 
19. Bowers, Michael A. 1987. Precipitation and the relative abundances of desert winter annuals: a 6-year study in the northern Mohave Desert. Journal of Arid Environments. 12: 141-149. 
20. Bowns, James E.; West, Neil E. 1976. Blackbrush (Coleogyne ramosissima Torr.) on southwestern Utah rangelands. Research Report 27. Logan, UT: Utah State University, Utah Agricultural Experiment Station. 27 p. 
21. Brooks, Matthew L. 1999. Alien annual grasses and fire in the Mojave Desert. Madrono. 46(1): 13-19. 
22. Brooks, Matthew Lamar. 1998. Ecology of a biological invasion: alien annual plants in the Mojave Desert. Riverside, CA: University of California. 186 p. Dissertation. 
23. Brown, David E.; Lowe, Charles H.; Hausler, Janet F. 1977. Southwestern riparian communities: their biotic importance and management in Arizona. In: Johnson, R. Roy; Jones, Dale A., tech. coords. Importance, preservation and management of riparian habitat: a symposium: Proceedings; 1977 July 9; Tucson, AZ. Gen. Tech. Rep. RM-43. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 201-211. 
24. Brown, James K.; Smith, Jane Kapler, eds. 2000. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech Rep. RMRS-GRT-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 257 p. 
25. California Exotic Pest Plant Council. 1999. Pest plants of greatest ecological concern, [Online]. Available: http://www.caleppc.org/info/plantlist.html [2001, February 12]. 
26. Cave, George Harold, III. 1982. Ecological effects of fire in the upper Sonoran Desert. Tempe, AZ: Arizona State University. 124 p. Thesis. 
27. Connor, J. Michael; Willoughby, Bob L. 1997. Effects of blue oak canopy on annual forage production. In: Pillsbury, Norman H.; Verner, Jared; Tietje, William D., technical coordinators. Proceedings of a symposium on oak woodlands: ecology, management, and urban interface issues; 1996 March 19-22; San Luis Obispo, CA. Gen. Tech. Rep. PSW-GTR-160. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 321-326. 
28. Corbineau, F.; Belaid, D.; Come, D. 1992. Dormancy of Bromus rubens L. seeds in relation to temperature, light and oxygen effects. Weed Research. 32(4): 303-310. 
29. 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. 
30. Dittberner, Phillip L.; Olson, Michael R. 1983. The plant information network (PIN) data base: Colorado, Montana, North Dakota, Utah, and Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 786 p. 
31. Eliason, Scott A; Allen, Edith B. 1997. Exotic grass competition in suppressing native shrubland re-establishment. Restoration Ecology. 5(3): 245-255. 
32. Evans, Raymond A.; Biswell, Harold H.; Palmquist, Debra E. 1987. Seed dispersal in Ceanothus cuneatus and C. leucodermis in a Sierran oak-woodland savanna. Madrono. 34(4): 283-293. 
33. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. 
34. Florence, Melanie. 1986. Plant succession on prescribed burn sites at Pinnacles National Monument. Fremontia. 14(3): 31-33. 
35. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. 
36. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. 
37. Gordon, Aaron; Sampson, Arthur W. 1939. Composition of common California foothill plants as a factor in range management. Bull. 627. Berkeley, CA: University of California, College of Agriculture, Agricultural Experiment Station. 95 p. 
38. Halvorson, William L.; Clark, Ronilee A. 1989. Vegetation and floristics of Pinnacles National Monument. Tech. Rep. No. 34. Davis, CA: University of California at Davis, Institute of Ecology, Cooperative National Park Resources Study Unit. 113 p. 
39. Hansen, Robert Bruce. 1986. The effect of fire and fire frequency on grassland species composition in California's Tulare Basin. Fresno, CA: California State University, Fresno. 133 p. Thesis. 
40. Heady, Harold F. 1977. Valley grassland. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 491-514. 
41. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. 
42. Horton, J. S.; Kraebel, C. J. 1955. Development of vegetation after fire in the chamise chaparral of southern California. Ecology. 36(2): 244-262. 
43. Humphrey, R. R. 1950. Arizona range resources: II. Yavapai County. Bull. 229. Tucson, AZ: University of Arizona, Agricultural Experiment Station. 55 p. 
44. Hunter, Richard. 1991. Bromus invasions on the Nevada Test Site: present status of Bromus rubens and Bromus tectorum with notes on their relationship to disturbance and altitude. The Great Basin Naturalist. 51(2): 176-182. 
45. Jackson, Louise E.; Roy, Jacques. 1986. Growth patterns of Mediterranean annual and perennial grasses under simulated rainfall regimes of southern France and California. Acta Ecologica. 7(21): 191-212. 
46. Jones, Stanley D.; Wipff, Joseph K.; Montgomery, Paul M. 1997. Vascular plants of Texas. Austin, TX: University of Texas Press. 404 p. 
47. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with: The Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. 
48. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 2 volumes]. Dissertation. 
49. Kearney, Thomas H.; Peebles, Robert H.; Howell, John Thomas; McClintock, Elizabeth. 1960. Arizona flora. 2d ed. Berkeley, CA: University of California Press. 1085 p. 
50. Keeley, Jon E. 1990. The California valley grassland. In: Schoenherr, Allan A., ed. Endangered plant communities of southern California: Proceedings of the 15th annual symposium; 1989 October 28; Fullerton, CA. Special Publication No. 3. Claremont, CA: Southern California Botanists: 2-23. 
51. Keeley, Jon E.; Keeley, Sterling C. 1984. Postfire recovery of California coastal sage scrub. The American Midland Naturalist. 111(1): 105-117. 
52. Keeley, Sterling C.; Keeley, Jon E.; Hutchinson, Steve M.; Johnson, Albert W. 1981. Postfire succession of the herbaceous flora in southern California chaparral. Ecology. 62(6): 1608-1621. 
53. Kirkpatrick, J. B.; Hutchinson, C. F. 1977. The community composition of Californian coastal sage scrub. Vegetatio. 35(1): 21-33. 
54. Kirkpatrick, J. B.; Hutchinson, C. F. 1980. The environmental relationships of Californian coastal sage scrub and some of its component communities and species. Journal of Biogeography. 7: 23-38. 
55. Krausman, Paul R.; Kuenzi, Amy J.; Etchberger, Richard C.; [and others]. 1997. Diets of mule deer. Journal of Range Management. 50(5): 513-522. 
56. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. 
57. Laude, Horton M. 1956. Germination of freshly harvested seed of some western range species. Journal of Range Management. 9: 126-129. 
58. Loftin, Samuel Robert. 1987. Postfire dynamics of a Sonoran Desert ecosystem. Tempe, AZ: Arizona State University. 97 p. Thesis. 
59. McClaran, Mitchel P.; Brady, Ward W. 1994. Arizona's diverse vegetation and contributions to plant ecology. Rangelands. 16(5): 208-217. 
60. McPherson, James K.; Muller, Cornelius H. 1969. Allelopathic effects of Adenostoma fasciculatum, "chamise", in the California chaparral. Ecological Monographs. 39(2): 177-198. 
61. Montgomery, Kenneth R. 1976. Regeneration of introduced species of Cistus (Cistaceae) after fire in southern California. Madrono. 23(8): 417-427. 
62. Nelson, Deanna R.; Harper, Kimball T. 1991. Site characteristics and habitat requirements of the endangered dwarf bear-claw poppy (Arctomecon humilis Coville, Papaveraceae). The Great Basin Naturalist. 51(2): 167-175. 
63. O'Leary, John F.; Westman, Walter E. 1988. Regional disturbance effects on herb succession patterns in coastal sage scrub. Journal of Biogeography. 15: 775-786. 
64. Parker, Karl G. 1975. Some important Utah range plants. Extension Service Bulletin EC-383. Logan, UT: Utah State University. 174 p. 
65. Patel, P. M.; Wallace, A. 1980. Effect of trace metal applications on emergence of Bromus rubens in desert soil. Journal of Plant Nutrition. 2(1&2): 65-66. 
66. Pavlick, Leon E. 1995. Bromus L. of North America. Victoria, BC: Royal British Columbia Museum. 160. 
67. Paysen, Timothy E.; Derby, Jeanine A.; Black, Hugh, Jr.; [and others]. 1980. A vegetation classification system applied to southern California. Gen. Tech. Rep. PSW-45. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 33 p. 
68. Phillips, Barbara G. 1992. Status of non-native plant species, Tonto National Monument, Arizona. Technical Report NPS/WRUA/NRTR-92/46. Tucson, AZ: The University of Arizona, School of Renewable Natural Resources, Cooperative National Park Resources Study Unit. 25 p. 
69. Preston, Kris P. 1993. Selection for sulfur dioxide and ozone tolerance in Bromus rubens along the south central coast of California. Annals of the Association of American Geographers. 83(1): 141-155. 
70. Rasmussen, Lars L.; Brotherson, Jack D. 1986. Habitat relationships of sandsage (Artemisia filifolia) in southern Utah. In: McArthur, E. Durant; Welch, Bruce L., compilers. Proceedings--symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9-13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 58-66. 
71. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
72. Rogers, Garry F.; Steele, Jeff. 1980. Sonoran Desert fire ecology. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 15-19. 
73. Seegmiller, Rick F.; Krausman, Paul R.; Brown, William H.; Whiting, Frank M. 1990. Nutritional composition of desert bighorn sheep forage in the Harquahala Mountains, Arizona. Desert Plants. 10(2): 87-90. 
74. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. 
75. St. John, Harold. 1973. List and summary of the flowering plants in the Hawaiian Islands. Hong Kong: Cathay Press Limited. 519 p. 
76. Stace, Clive. 1997. New flora of the British Isles. 2nd ed. Cambridge, UK: The Bath Press, Inc. 1130 p. 
77. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
78. Thomas, Timothy W. 1987. Population structure of the valley oak in the Santa Monica Mountains National Recreation Area. In: Plumb, Timothy R.; Pillsbury, Norman H., technical coordinators. Proceedings of the symposium on multiple-use management of California's hardwood resources; 1986 November 12-14; San Luis Obispo, CA. Gen. Tech. Rep. PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 335-340. 
79. Turkowski, Frank J. 1975. Dietary adaptability of the desert cottontail. Journal of Wildlife Management. 39(4): 748-756. 
80. Tutin, T. G.; Heywood, V. H.; Burges, N. A.; [and others]. 1980. Flora Europaea. Vol. 5: Alismataceae to Orchidaceae (Monocotyledones). Cambridge, UK: University Press. 452 p. 
81. U.S. Department of Agriculture, Soil Conservation Service. 1994. Plants of the U.S.--alphabetical listing. Washington, DC: U.S. Department of Agriculture, Soil Conservation Service. 954 p. 
82. Utah State University. 2002. Grass manual on the web, [Online]. In: Manual of grasses for North America--Intermountain herbarium. Logan, UT: Utah State University (Producer). Available: http://herbarium.usu.edu/grassmanual/ [2006, January 10]. 
83. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. 
84. Wilken, Dieter. 2001. [E-mail to Janet Howard]. January 8. Santa Barbara, CA: Santa Barbara Botanic Garden. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab, Missoula, MT; RWU 4403 files. 
85. Wu, K. K.; Jain, S. K. 1978. Genetic and plastic responses in geographic differentiation of Bromus rubens populations. Canadian Journal of Botany. 56: 873-879. 
86. Wu, K. K.; Jain, S. K. 1979. Population regulation in Bromus rubens and B. mollis: life cycle components and competition. Oecologia. 39(3): 337-357. 
87. Zedler, Paul H.; Gautier, Clayton R.; McMaster, Gregory S. 1983. Vegetation change in response to extreme events: the effect of a short interval between fires in California chaparral and coastal scrub. Ecology. 64(4): 809-818.