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FEDERAL LEGAL STATUS:
DISTRIBUTION AND OCCURRENCE
Blackbrush occurs primarily in the transition zone between the Mojave and Great Basin deserts and on the western border of the Sonoran Desert, forming a band from southeastern California to southwestern Colorado. It also occurs along the borders between Nevada, Utah, and Arizona [2,6,17], predominantly along the Colorado and San Juan river drainages in southeastern Utah and in adjacent drainages in the Mojave and Great Basin transition [6,8,17]. Blackbrush has also been reported as an invasive brush species in western, west-central, and southwestern Texas grasslands .
KUCHLER  PLANT ASSOCIATIONS:
SRM (RANGELAND) COVER TYPES :
HABITAT TYPES AND PLANT COMMUNITIES:
In California, blackbrush occurs as a subdominant species in the Mojave mixed woody scrub, and as a common understory species in pinyon-juniper (Pinus-Juniperus spp.) woodlands . It is characteristic of the northern desert scrub in Arizona [32,44], and is dominant in the canyon desert areas of northern Arizona, comprising 90% of the vegetation in areas where it occurs . Blackbrush is important in the Colorado pinyon (P. edulis)/blackbrush habitat type in northern Arizona . In Nevada, blackbrush forms a dominant association [106,125], is a primary shrub associate in the sagebrush association and the salt desert scrub association, and is a common associate in the creosotebush-white bursage, hopsage (Grayia spinosa), and Mojave mixed scrub associations . It is a subdominant in pinyon-juniper woodlands [4,5,13,21,103,106] and a codominant in stands of Joshua tree [1,61,77,103] and shadscale . It occurs as a dominant shrub in both pinyon-dominated and juniper-dominated woodlands, occurring in the following plant associations :
northern Mojave: singleleaf pinyon (P.
monophylla)/blackbrush/blue grama (Bouteloua gracilis); singleleaf
pinyon/blackbrush/wavyleaf Indian paintbrush (Castilleja
applegatei); singleleaf pinyon/blackbrush/ Sandberg bluegrass (Poa secunda);
Utah juniper (J. osteosperma)/blackbrush/blue grama,
Utah juniper/blackbrush/Indian ricegrass (Achnatherum hymenoides); Utah juniper/blackbrush/mutton grass (Poa fendleriana)
In pinyon-juniper woodlands blackbrush is commonly associated with Joshua tree, Our Lord's candle (Y. whipplei), sagebrush, ephedra (Ephedra spp.), winterfat, and cactus (Opuntia spp.) [55,56,100,105,126].
Relative blackbrush density varies dramatically by plant community type; for example, 8,894 plants /ha were found in the blackbrush scrub community type, while 647 plants/ha and 0 plants/ha were found on Joshua tree woodland and Mojave mixed steppe sites, respectively . In 1 canyon in California, blackbrush cover ranged from 2.2% to 20.6% of total plant cover . In monospecific blackbrush shrublands, plant density and species diversity are much lower than in adjacent shrublands [36,54]. On blackbrush sites in Arizona, blackbrush contributed 82% to 95% of shrub cover . Though few other shrubs occur in these stands, red brome (Bromus madritensis ssp. rubens) and cheatgrass (B. tectorum) may be the dominant understory species [16,17,18,23]. At sites in southwestern Utah, blackbrush contributed 75% of all plant cover, with red brome and cheatgrass accounting for 12% . Big galleta (Pleuraphis rigida) also contributes substantially to understory vegetation in blackbrush communities . The blackbrush overstory tends to preclude extensive understory development [19,125]; however, herbaceous plants tend to occur in greatest abundance on the periphery of blackbrush canopies, indicating the presence of a more favorable microenvironment near the shrubs and a lack of any toxic effect exerted by blackbrush [17,23].
Blackbrush also commonly occurs with the following shrubs and grasses: fourwing saltbush (Atriplex canescens), desert bitterbrush (Purshia glandulosa), shrub live oak (Quercus turbinella), Cooper's heathgoldenrod (Ericameria cooperi), Fremont's dalea (Psorothamnus fremontii), white burrobush (Hymenoclea salsola), Anderson wolfberry (Lycium andersonii), broom snakeweed (Gutierrezia sarothrae), desert globemallow (Sphaeralcea ambigua), spiny mendora (Menodora spinescens), bladdersage (Salazaria mexicana), blue yucca, banana yucca (Y. baccata), galleta (Pleuraphis jamesii), threeawn (Aristida spp.), arid needlegrass (Achnatherum arida), desert needlegrass (A. speciosum), needle-and-thread grass (Hesperostipa comata), black grama (Bouteloua eriopoda), blue grama (B. gracilis), western wheatgrass (Pascopyrum smithii), and California buckwheat (Eriogonum fasiculatum) [1,5,13,16,62,65,69,71,89,95,96,107,118,119,124].
VALUE AND USE
Blackbrush is not preferred as forage by domestic livestock, deer, or pronghorn, but it does provide poor forage during the spring, summer, and fall for domestic cattle, horses, and domestic sheep [8,19,41,50,90,112,124]. Blackbrush provides poor to good forage for domestic goats [41,84,90]. Mule deer and bighorn sheep generally use the blackbrush vegetation type in winter, and livestock generally use it in winter and spring [17,98,112,124]. The principal forage value of blackbrush appears to be as a browse species for bighorn sheep [17,18,68]; blackbrush communities also provide important habitat for desert bighorn sheep in Nevada . Domestic sheep and goats, and to a lesser extent, cattle browse blackbrush [47,70,118,124]. It provides fair forage for domestic sheep and cattle in the winter in southwestern Utah [17,41], but spiny stems coupled with chemical compounds in current year's growth protect blackbrush from heavy browsing by livestock [6,79,80]. Due to the spinescent growth form of blackbrush, the low amounts of current-season growth are of limited accessibility to most browsing animals [79,112]. Carrying capacities on blackbrush ranges are low, from 30 to 300 acres/AUM [8,124].
Blackbrush often occurs as nearly monospecific stands, with few other forage species available . Though it is not desirable deer forage, in areas where it is extensive it may experience heavy browsing pressure [8,68]. Where it forms an appreciable proportion of the brush cover, it provides a substantial part of the diet of domestic sheep and goats and deer, despite its small leaves and spinescent growth . In California, blackbrush has comprised up to 25% of the mule deer winter diet . Blackbrush-dominated sites where animals congregate may provide inadequate nutrition and impede the regeneration of bitterbrush, a key browse species . The presence of blackbrush in Arizona chaparral may reduce the range quality rating due to its low forage value and exclusion of higher forage value plants . Greater resistance to grazing probably allows blackbrush to persist and perhaps expand after the more palatable species are removed .
The spinescent character of blackbrush combined with low phosphorus and protein levels and high tannin levels result in low palatability [8,80]. New growth is likely more palatable and nutritious than old spinescent branches , but regrowth of blackbrush has been found to be unpalatable to domestic goats and cattle .
Blackbrush is low in phosphorus and protein and high in poorly digestible fiber [8,79,84,85]. Low nitrogen levels suppress microbial activity in the rumen, thereby decreasing fiber digestibility and lowering forage intake. High tannin levels, also typical of blackbrush, may depress intake by decreasing palatability and suppressing protein digestion . Domestic goats have been found to avoid blackbrush current-season growth in favor of older growth, even though new growth is higher in nitrogen and is more digestible. Current-season growth is high in concentrated tannins, which may deter browsing because tannins interfere with digestive or metabolic processes .
Twig nutrition varies by location on the plant; twigs
from basal branches, located within the canopy, are higher in crude protein and
in-vitro digestible matter than those on older, terminal branches, located at
the outer edges of the canopy [79,82,86]. Blackbrush nutrition has been evaluated according to plant part
[16,17,118]: the following table compares
the nutritional content of leaves and stems [16,17]:
Blackbrush leaves and stems exceed the minimum carotene required for gestating and lactating domestic animals but are deficient in phosphorus for domestic cattle and sheep during gestation and lactation [16,17]. Ether extract is comparable to that of big sagebrush and black sagebrush (Artemisia nova) during the winter [16,17].
VALUE FOR REHABILITATION OF DISTURBED SITES:
Blackbrush displays no natural vegetative reproduction , but regeneration can be achieved with asexual propagation from artificial cuttings. One-year growth has been found to produce a higher percentage of rooted cuttings, more roots, and longer roots than older growth .
OTHER USES AND VALUES:
Removal of spinescent material from blackbrush plants stimulates sprouting from basal and axillary buds; therefore, plants that are heavily browsed by livestock produce large quantities of new, more accessible growth [79,80,85,112]. Browsing improves nutritional quality of blackbrush twigs by increasing current-season growth; however, browsing may decrease palatability due to high tannin levels in current-season growth . Nutritional value varies in response to browsing treatments: low protein and high tannin content persists, though current-season growth generally has increased protein . Heavy browsing followed by 1 to 2 years of rest allows blackbrush to accumulate twigs that are more palatable because they are lower in tannins due to lower proportions of current-season growth [80,84,85,86]. Domestic goats can be used to remove spinescent growth and increase production of current-season growth to improve forage for cattle; however, the low crude protein levels may cause a reduction in livestock weight [83,112]. Livestock browsing on blackbrush should be supplemented with protein to improve rumen function and minimize weight loss [80,84,85]. With intensive management, stocking intensities of 1.8 AUM/ha can be maintained .
Blackbrush can produce substantially larger amounts of current growth with somewhat greater palatability and nutritional value through the use of mechanical or browsing treatments . Prescribed burning can lead to replacement by other shrubs, diversifying plant communities and increasing the winter forage base, which may increase livestock carrying capacity [8,112]. Because blackbrush exhibits strong apical dominance that suppresses annual twig growth, removal of terminal buds during the dormant winter season stimulates lateral twig growth during the spring [81,82]. Brush beating damages plants and stimulates growth of new shoots, improving forage quality . The results of a simulated brush beating in different plant communities 1 year after treatment are presented below. Blackbrush plants in the blackbrush association responded better to the brush beating treatments than blackbrush plants in either the Joshua tree/blackbrush or Utah juniper/blackbrush associations. The number of blackbrush plants (out of 30 at each location) is presented according to the response to brush-beating treatment and the plant community in which the blackbrush occurred :
Efforts to manipulate blackbrush rangelands to increase forage production have produced unanticipated results . Blackbrush stands might be manipulated for improvement of forage quality and quantity without destroying the original vegetation; however, the manipulation may open the plant community to the invasion of other, perhaps less desirable, species .
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
Blackbrush is a native, aromatic shrub with soft wood , growing from 1 to 6 feet (0.3-2 m) tall [16,27,38,79,84,85,90,117,123]. It shows compact, erect growth, with a symmetrically round form [16,118]. The scientific name refers to the unusual sheath or torus around the ovary (Coleogyne) and to its many-branched morphology (ramosissima) . The common name is derived from the color of the dense branches, which have gray bark that turns black with age or when wet [8,16,17,90]. The terminal branches grow for a few years then die, drying back for several centimeters from the tip and resulting in the characteristically tangled spinescence of blackbrush [16,17,79,84,85]. Apical dominance is removed when the terminal buds die, allowing development of lateral branches [16,78,79,84,85]. The shrub undergoes stem-splitting, in which the main stem splits into several smaller portions [16,17]. These clusters of multi-stemmed segments also correspond to separate segments of the root system .
Blackbrush has a diffuse and shallow root system . The greatest root biomass of blackbrush is found at a soil depth of 4 to 12 inches (10-30 cm), with few roots penetrating the fractured caliche layer, if present [16,79,84,85]. Large supporting roots are located directly beneath the plant, and root biomass tends to decrease with increasing distance from the plant and with increasing soil depth [79,84,85]. Shallow soils often result in a low root:shoot ratio and limited root development in blackbrush communities .
Blackbrush is evergreen [1,27,79,84,85,89,94,109], though it may lose substantial leaf area during the dry summer season . Blackbrush is drought-deciduous, avoiding water stress by becoming temporarily dormant and shedding older leaves as stress intensifies during the dry season . After leaf drop, it enters a long summer dormancy 
Atypical of the rose family [8,17], blackbrush flowers typically lack petals [38,68,90,123]. Blackbrush flowers are perfect, solitary, and terminal on the young branchlets . The fruits are dry, leathery achenes, 3 to 4 mm long, with a bent and twisted style [38,90,117].
Blackbrush has a "long" life span [92,121], and its life history emphasizes maintenance of existing individuals; establishment from seed is rare . Blackbrush-dominated stands are generally monotypic, simple communities where shrub cover is high. Close spacing permits little growth of other vegetation [8,16,17,41,45,58,79,84,85,94].
Blackbrush generally has a low germination rate [9,121], but with heavy, early spring rains, blackbrush seeds germinate in relatively large numbers. These may be the only conditions under which substantial germination occurs, suggesting that "pulse" climatic events are needed for establishment [9,10,121]. Soil moisture is required before seeds will break dormancy; watering at 2-week intervals was found to increase germination more than watering at 1- or 3-week intervals . Germination of blackbrush seeds requires cold stratification without light [16,17,53]. Germination has been found to increase from 53% with no treatment to 83% after 7 weeks of cold stratification at 39 degrees Fahrenheit (4 oC) [16,17,29,53]. Seeds have also been found to respond to a moist storage at 41 degrees Fahrenheit (5 oC) and germinate at that temperature .
Germination patterns vary as a function of climate and elevation. Seeds collected from low-elevation sites were less dormant than seeds from high elevation sites in southern Utah and Nevada; the seeds from the 3,930-foot (1200 m) sites required a shorter chilling period to increase germination response than those from the 5,085-foot (1550 m) sites . This relationship between dormancy status and site elevation may indicate that blackbrush has evolved ecotypes [53,75]. Dreeson and Harrington  found that substantial age and source differences are apparent in regard to population germination and/or stratification requirements. The sensitivity of seeds to salinity may be a limiting factor governing the distribution of blackbrush [17,118].Blackbrush has a slow growth rate [57,121] that may be the result of shallow soils and an often-present caliche layer, which impede root growth and soil moisture .
Blackbrush stands occur on well-drained sites including alluvial and colluvial slopes, washes, valley bottoms, lowlands, and flatlands of mild slope, and derived from limestone, sandstone, gneiss, and basalt [1,2,5,6,8,10,15,16,23,47,51,55,58,106,107,118,123,124]. Soils supporting blackbrush are generally shallow, poorly developed, and coarse textured, often with abundant exposed rock and high sand content [5,6,8,17,23,44,51,98,118,123]. These sites are also calcareous, moderately alkaline, and low in salinity [9,17,94,118] with pH ranging from 7.8 to 8.5 [16,17,23]. Blackbrush has a low tolerance for salinity, excessive soil moisture, and impeded soil aeration . Shrubs are often clustered on small mounds, evidently created by entrapment of wind-blown material . There is typically a well-developed microphytic crust on the soil surface between shrubs .
The shallowness of soil may in part determine the abundance and distribution of blackbrush [23,55]. Blackbrush is abundant on shallow soils with caliche layers [2,8,12,16,17,23,55,118,125], but is more abundant on adjacent, deeper soils . Blackbrush occurs on ancient granitic debris flows in California, with the cover and density of blackbrush increasing with the age of the debris [120,121]. On the oldest depositional area, blackbrush is nearly monospecific, and the only physical differences between the flows studied are those resulting from difference in geologic age; notably, the intermediate and oldest flows had caliche layers [120,121]. The association of blackbrush with old soils and the lack of it on young basalt flows implies either an allogenically controlled succession with blackbrush dominating  or an affinity of blackbrush for calcium carbonate irrespective of the age of the surface [121,125].
Blackbrush individuals alter the soil chemistry around their bases [16,17,121]. Bowns  found that percent totals of nitrogen and available phosphorus are higher in soil beneath blackbrush plants than in the spaces between, and both nutrients decrease with increasing soil depth .
Blackbrush flowers only in spring, probably the result of photoperiodism . Flowering is induced by moderate to heavy winter precipitation [9,56]. Increases in winter precipitation and resulting soil moisture contribute to an abundance of flowers and seeds . Flower buds begin to develop at the tips of terminal or lateral branches 2 weeks after the shoot growth begins and are fully open after 5 weeks [1,16,17]. By 6 weeks, 80 to 100% of the flowers are open, and no further twig elongation occurs once the flowers are fully developed [16,17]. Blackbrush at high elevations has a shorter flowering period than blackbrush at lower elevations, and flowering on individual plants is not synchronous, occurring over a 1- to 3-week period . Fruits begin to develop in late April and early May [1,16,17].
After growth ceases in June, older outermost leaves yellow and dry out. Abscission occurs in July and August, causing a large buildup of organic matter [1,2,16,17]. Blackbrush may lose most of its leaves during summer dormancy, but retains enough leaves at the ends of branches to be considered an evergreen species [1,2].
Blackbrush phenophases vary according to location within its range. The following table lists the mean initial date of each phenophase along an elevation gradient from 4,900 to 5,900 feet (1,500-1,800 m) in southern Nevada. Standard errors and significant ( p£0.05) differences are denoted by a and b, respectively .
Blackbrush stands are subject to fire , and fire will start and spread easily due to the dense, close spacing, "tinder-like" nature, and resinous foliage of blackbrush [17,116,124,125]. There are usually few forbs or grasses in blackbrush stands that might aid in carrying fire, but despite this, blackbrush communities burn under conditions of high temperature, high wind velocity, and low relative humidity [42,72]. Fire also occurs in blackbrush stands on sites with high proportions of herbaceous perennial species or in years in which annuals are abundant . Blackbrush's often strong association with red brome and cheatgrass may result in higher fire frequencies than would occur without the bromes: the grasses leave dense, persistent dead stems that promote fire spread . Various sprouting shrubs and annuals establish after fire, and once these species gain dominance the recurrence rate for fire increases . The presence of Joshua trees in blackbrush stands may also contribute to increased fire frequencies due to lightning strikes on the Joshua trees .
Because blackbrush is a nonsprouter, very susceptible to fire, and slow to reinvade sites, it is removed by fire [17,128], and succeeding communities are variable . Fire in blackbrush stands in southwestern Utah resulted in a variety of species dominating the postfire vegetation [3,17]. These postfire dominants include turpentine bush, desert bitterbrush, desert almond, big sagebrush, and some nearly monospecific stands of broom snakeweed . Grasses are more abundant in burned blackbrush communities , and burning may improve forage productivity . One 10-year-old burn in blackbrush was devoid of blackbrush and dominated by brittlebrush and desert mallow, with a denser cover of red brome and cheatgrass compared to exotic brome cover in the adjacent unburned blackbrush community . In another study of burned blackbrush sites in southwestern Utah, most shrubs were removed by fire. In the 1st postfire year, forbs greatly increased and grasses moderately increased. Forbs steadily decreased over time, approaching prefire levels, while grasses steadily increased, peaked at postfire year 6, and then declined to prefire levels. Shrub dominance on these sites returned within 20 years, but shrub composition after burning only slightly resembled composition before fire. Blackbrush cover was greatly reduced on all sites. Cryptogamic soil crusts associated with blackbrush communities were also strongly affected by fire. Before burning cryptogamic crusts contributed 9% of plant cover but were reduced to less than 1% of total plant cover after fire. There was very little evidence of crust formation after 19 postburn years . Fire has promoted succession to grassland by destroying the cryptogamic crust, which stabilizes the soil .
Blackbrush fire regime: The blackbrush association is composed of dense to scattered low-stature shrubs and dense to open grasses, and it maintains the highest cover of any desert shrub community. Blackbrush experiences a stand-replacement fire regime, though historical documentation of blackbrush fire cycles is limited. Frequent large fires have eliminated blackbrush from some areas. Fuel production in blackbrush ranges from 250 to 500 lbs/acre, and blackbrush is negatively associated with fine fuels of litter and grasses. Blackbrush occurs in areas with approximately 7 inches (180 mm) of annual precipitation, and cyclic desert precipitation above 10 to 14 inches (250-360 mm) may increase biomass and fuel continuity enough to increase fire behavior potential .
The following table provides some fire regime intervals for ecosystems where blackbrush occurs:
Blackbrush stands are substantially decreased or eliminated by fire ; fire usually kills blackbrush seeds and mature shrubs [17,22]. Blackbrush is susceptible to fire and slow to reestablish ; it is generally removed from the site for 25 to 30 years [8,129].
PLANT RESPONSE TO FIRE:
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
FIRE MANAGEMENT CONSIDERATIONS:
Fires may increase species diversity, livestock carrying capacity, and range condition. The following table describes 5 different sites and improvement of range condition after fire. All sites had cobbled, stony soils with 35% or more gravel content and received a mean annual precipitation of 125 mm .
Prescribed burning on 3 southern Nevada sites killed the blackbrush cover, and the species failed to reestablish after as long as 28 years. Plant succession varied widely, with different plant species dominating on different burns, but the density of annual species was substantially increased in the 1st 3 years following burning. Replacement shrubs after fire were largely undesirable forages . Since sites may be highly suitable for blackbrush, burning these areas to convert them to grassland may give unpredictable or undesirable results . After burning on 3 proximal sites in southwestern Utah, 1 site was dominated by turpentine bush, desert bitterbrush, desert almond, and big sagebrush; another site established a pure stand of broom snakeweed, and a 3rd site established a pure stand of big sagebrush . Vast areas of blackbrush in Nevada were burned in the 1940s and 1950s. These sites were subsequently occupied by annuals and broom snakeweed and have been subject to recurring fires. In wet years, the burned areas had 8 to 10 times more herbage production than the range before it was burned, but herbs were very sparse in dry years . Site potential is an important consideration for burning blackbrush; fire may be more useful on areas with better-developed soils and potential to revegetate to more desirable plants . Widespread burning to reduce blackbrush is not recommended due to the unpredictability of successive vegetation, accelerated soil erosion, long-term or permanent removal of blackbrush, and damage to cryptogamic soil crusts [24,125,129].
Fire may be a necessary tool to modify fuel buildup; however, research is needed regarding management and restoration recommendations for blackbrush .
Coleogyne ramosissima: References
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29. Dreesen, David R.; Harrington, John T. 1997. Propagation of native plants for restoration projects in the southwestern U.S.--preliminary investigations. In: Landis, Thomas D.; Thompson, Jan R., tech. coords. Regeneration, reforestation, restoration: The seedling is the key; National proceedings, forest and conservation nursery associations--1997; Gen. Tech. Rep. PNW-GTR-419. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 77-88. 
30. Everett, Richard L. 1985. Great Basin pinyon and juniper communities and their response to management. In: Symposium on the cultural, physical and biological characteristics of range livestock industry in the Great Basin: Proceedings, 38th annual meeting of the Society for Range Management; 1985 February 11-14; Salt Lake City, UT. Denver, CO: Society for Range Management: 53-62. 
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Coleogyne ramosissima Index
Related categories for SPECIES: Coleogyne ramosissima | Blackbrush
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