Photo courtesy of Pat Breen, Oregon State University
Black oak (Quercus velutina)/greenleaf manzanita in the hardwood rangelands 
Whitethorn ceanothus (Ceanothus cordulatus)-greenleaf manzanita-bush chinquapin (Chrysolepis sempervirens) in the San Bernardino Mountains 
Jeffrey pine (Pinus jeffreyi)/greenleaf manzanita-snowbrush ceanothus (Ceanothus velutinus), found throughout the Sierra Nevada at mid- to upper elevations (5,920-9,520 feet (1,800-2,900 m)) 
Canyon live oak (Q. chrysolepis)/whiteleaf manzanita-greenleaf manzanita in Castle Crags State Park, Shasta County
Wedgeleaf ceanothus-California coffeeberry (C. cuneatus-Rhamnus californica)-greenleaf manzanita in Castle Crags State Park 
Pacific ponderosa pine (Pinus ponderosa var. ponderosa)/greenleaf manzanita in Lava Beds National Monument
Pacific ponderosa pine/antelope bitterbrush (Purshia tridentata)-greenleaf manzanita near the extreme southern border of Lava Beds National Monument
Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita/western needlegrass (Achnatherum occidentale ssp. occidentale) at the highest elevations of Lava Beds National Monument 
Singleleaf pinyon/pale serviceberry (Pinus monophylla-Amelanchier pallida)/ greenleaf manzanita in the eastern part of Mathews Canyon Watershed 
Greenleaf manzanita/snowbrush ceanothus-prostrate ceanothus (C. prostratus) shrublands 
Singleleaf pinyon/Saskatoon serviceberry (A. alnifolia)/greenleaf manzanita woodlands 
Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita community in the Cascade Range of south-central Oregon 
Tanoak (Lithocarpus densiflorus)/manzanita (greenleaf, whiteleaf, and hairy manzanita (Arctostaphylos columbiana)/beargrass (Xerophyllum tenax) east of the Coast Ranges crest in the southwestern portion of the state from approximately 3,000 to 3,700 feet (900-1,100 m) 
Pacific ponderosa pine/greenleaf manzanita/Idaho fescue (Festuca idahoensis) on the Silver Lake mule deer range from 4,900 to 6,500 feet (1,500-2,000 m)  and in other south-central portions of the state 
Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita on pumice soils in the south-central portion of the state 
Greenleaf manzanita/snowbrush ceanothus in the eastern Siskiyou Mountains 
White fir-Pacific ponderosa pine/greenleaf manzanita-pinemat manzanita-Oregon-grape (A. nevadensis-Berberis repens) in the Fremont-Winema National Forest from 5,000 to 6,500 feet (1,500-2,000 m) 
Sierra lodgepole pine (Pinus contorta var. murrayana)/snowbrush ceanothus-greenleaf manzanita-pinemat manzanita in the Deschutes and Fremont-Winema National Forests from 4,800 to 6,000 feet (1,500-2,000 m)
Sierra lodgepole pine/pinemat manzanita-greenleaf manzanita in the Deschutes and Fremont-Winema National Forests from 5,800 to 7,000 feet (1,800-2,000 m)
Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita/Columbia needlegrass (Achnatherum nelsonii) in the Deschutes and Fremont-Winema National Forests from 3,000 to 5,900 feet (900-1,800 m)
Pacific ponderosa pine-white fir-sugar pine (Pinus lambertiana)/snowbrush ceanothus-greenleaf manzanita in the Deschutes and Fremont-Winema National Forests from 4,100 to 5,900 feet (1,200-1,800 m)
Pacific ponderosa pine-Sierra lodgepole pine-white fir/greenleaf manzanita-snowbrush ceanothus/yellow sedge-glaucous beardtongue (Penstemon euglaucus) in the Deschutes National Forest from 4,300 to 5,600 feet (1,300-1,700 m)
California red fir (Abies magnifica)-white fir-western white pine (Pinus monticola)/pinemat manzanita-greenleaf manzanita in the Deschutes and Fremont-Winema National Forests from 5,450 to 7,000 feet (1,660-2,000 m) 
Interior ponderosa pine (Pinus ponderosa var. scopulorum)/Idaho fescue (greenleaf manzanita phase) in the south-central Uinta Mountains; this habitat type is associated with warm, dry sites 
Interior ponderosa pine/greenleaf manzanita in Garfield County and east to the LaSal and Abajo mountains 
Interior ponderosa pine/greenleaf manzanita in the Rocky Mountain and Intermountain West regions of southern Utah and western Colorado 
Interior ponderosa pine/Idaho fescue greenleaf manzanita phase in the Rocky Mountain and Intermountain West regions of eastern Washington, Idaho, northern Utah, central and southeastern Montana, and north-central Wyoming from 2,500 to 6,000 feet (760-2,000 m), and south-central Colorado from 8,800 to 9,300 feet (2,700-2,800 m) 
Photo courtesy of Pat Breen, Oregon State University
Greenleaf manzanita is an erect shrub with stout branches [94,106] that grows from 3 to 7 feet (1-2 m) tall [47,70,90,92,94,106,160]. Individual plants may have an ultimate spread of 10 feet (3 m) . Spreading or drooping leaves (never vertical) [89,92,160] are 0.8 to 2 inches (2-6 cm) long, 0.6 to 2 inches (1.5-4 cm) broad [90,92,94,160,226], and produced on twisted, crooked, or gnarled stems . The flowers, about 6 mm long , are borne on condensed, many-flowered panicles at the ends of some of the branches [94,167]. The fruit is a drupe, roughly 7-10 mm broad [90,92,160]. Each drupe contains approximately 5 stony, 1-seeded nutlets [70,120].
Underground parts: Unlike greenleaf manzanita populations elsewhere, greenleaf manzanita in the Sierra Nevada and southwestern Oregon has a lignotuber [1,89,94]. The lignotuber is described as a heavy, turnip- or globular-shaped organ that may form tabular platforms [92,99,102]. Greenleaf manzanita roots grow deep into the ground [119,197]. In the southern Sierra Nevada, greenleaf manzanita roots extend to a depth of no greater than 7 feet (2 m). Roots in the latter 5.74 feet (1.74 m) grew down through weathered, porous granitic bedrock . The roots of greenleaf manzanita are commonly infected by ectomycorrhizae or arbutoid mycorrhizae in the Sierra Nevada .
Physiological characteristics: Greenleaf manzanita is a drought-tolerant shrub . DeLucia and Schlesinger , based on data collected from Alpine County near Reno, Nevada, describe greenleaf manzanita as having high drought tolerance and nitrogen-use efficiency, but low water-use efficiency. Greenleaf manzanita is adapted to high levels of water stress. Greenleaf manzanita plants in the Sierra Nevada maintained a xylem potential of >-0.7 MPa throughout the growing season even during periods of drought .RAUNKIAER  LIFE FORM:
Pollination: Greenleaf manzanita is insect pollinated .
Breeding system: The mating system of greenleaf manzanita is primarily outcrossing .
Seed production: Greenleaf manzanita produces large seed crops nearly every year . In a mixed-conifer community of the Sierra Nevada, greenleaf manzanita populations produced a mean of 10,000 seeds/acre . In general, greenleaf manzanita plants do not begin to flower and fruit until the age of 8 to 10 . Where greenleaf manzanita populations sprout from a lignotuber, seed production is less than in nonsprouting populations [98,120].
Seed dispersal: Greenleaf manzanita seeds are dispersed by birds [139,175] and mammals including rodents, American black bears, and coyotes [112,117,139,175]. Seeds are dispersed from late summer  until the following spring [139,175]. The vast majority of Arctostaphylos spp. seeds are dispersed beneath the parent's canopy .
Seed banking: Greenleaf manzanita utilizes a seed bank [143,171,202]. The hard seeds of greenleaf manzanita can remain dormant in the soil for hundreds of years until stimulated to germinate [50,108].
Germination: Greenleaf manzanita seeds require scarification (by heat or disturbance such as logging activities) [67,99,109] followed by a period of cold stratification [18,109] for germination to occur. Seeds generally germinate on burned sites in the spring . Seedling densities as high as 25,233/ha have been reported following fire in the northern Sierra Nevada .
Fire-induced germination: Seeds can be stimulated to germinate by heat and/or chamise (Adenostoma fasciculatum) charate, followed by stratification [116,117]. Keeley  investigated the effects light, darkness, temperature, and charate had on greenleaf manzanita seed germination. Germination trials were conducted on filter paper and in soil. Greenleaf manzanita seeds germinated significantly (P<0.001) better in the filter paper treatment, and the following results are from the seeds germinated on filter paper. Temperature treatments were an unheated control, 158 °F (70 °C) for 1 hour; 212 °F (100 °C) for 5 minutes; and 248 °F (120 °C) for 5 minutes. One set of seeds received a 0.50-g application of chamise charate prior to temperature treatments. Following charate and temperature treatments, the seeds were stratified for 1 month at 41 °F (5 °C) and then incubated in the light or dark treatment for 3 weeks at 73 °F (23 °C). Light had a significant (P<0.001) negative effect on greenleaf manzanita germination. Further, in the light treatment, seeds subjected to 248 °F for 5 minutes germinated significantly (P<0.05) better than either seeds exposed to 158 °F for 1 hour or the no-heat control seeds. In the dark treatment, seeds exposed to charate and either no heat or 212 °F for 5 minutes germinated significantly (P<0.05) better than seeds not exposed to charate and either no-heat or 212 °F treatments. Further, in the dark/charate treatments, greenleaf manzanita seed germination significantly (P<0.05) decreased in the 248 °F treatment .
Greenleaf manzanita germination with light, heat, and charate treatments
Seedling establishment/growth: Greenleaf manzanita seedlings rarely occur on unburned or undisturbed sites . James  states that "fire prepares a soil environment conducive to greenleaf manzanita seedling growth by increasing both the ash layer and soil pH, and destroying allelopathic soil chemicals". Seedling establishment generally occurs the first postfire year . In a montane chaparral site in northern California, greenleaf manzanita seedlings establishing after fire reached a height of 3 feet (1 m) within 3 years .
In the first few years of development, greenleaf manzanita seedlings generally grow multiple stems. In May 1992, greenleaf manzanita seedlings ranging from 2 to 4 inches (5-10 cm) tall were planted at an elevation of 3,600 feet (1,100 m) on an unburned site in the Shasta-Trinity National Forest, California. Greenleaf manzanita seedlings were single stemmed at the time of planting and remained so throughout the first growing season. New sprouts appeared by the second growing season. At the end of the second growing season, 76% of seedlings had multiple stems, and that proportion remained constant throughout the 4-year study. At the end of the 1995 growing season, 89% of the seedlings had survived, yet none had produced flowers .
|Mean growth of greenleaf manzanita seedlings at the end of the 1992-1995 growing seasons|
|Crown width (cm)||10.9||27.8||47.7||68.6|
Vegetative regeneration: Greenleaf manzanita regenerates vegetatively by layering [24,90,94,98,106,113,140,141] and sprouting from the lignotuber [59,62,76,80,93,94,98,99,149,188,224,225].
Greenleaf manzanitas with lignotubers sprout following total or partial top-kill. In an early May clipping study in the Siskiyou Mountains in southwestern Oregon, greenleaf manzanita was either cut to ground level or had 50% of the plant removed. While plants cut to ground level did not show immediate growth, plants with growth only partially removed sprouted by 1 July. By posttreatment year 1, greenleaf manzanita density was substantially greater after clipping than before treatment on both sites .
Mean density and basal area of greenleaf manzanita plants subjected for total and partial top-kill
|Stems/ha||Basal area (m²/ha)|
|1 year after treatment||308,504||1.40|
|1 year after treatment||208,098||4.72|
Climate: Greenleaf manzanita is well adapted to hot, dry climates and can withstand wide temperature extremes . In the Sierra Nevada, where greenleaf manzanita is prevalent, the climate is characterized by hot, dry summers and mild, moist winters. Most of the precipitation occurs as snow from November to May [165,181,188].Elevation:
|Greenleaf manzanita elevational range|
|Arizona||7,000 to 8,500 feet |
|California||2,000 to 11,000 feet [90,160,183]|
|Colorado||7,000 to 9,000 feet [88,119]|
|Nevada||2,000 to 9,000 feet [106,192]|
|Utah||4,500 to 9,300 feet [167,226]|
|Sierra Nevada montane chaparral||5,000 to 7,000 feet |
Environmental conditions: Greenleaf manzanita is well adapted to high foliar temperatures and low soil moisture availability .
Slope and aspect: Greenleaf manzanita is most common on south and southwestern aspects where full sunlight is available [59,95].
Soil: Greenleaf manzanita typically occurs on soils that are well drained [96,119,149], shallow to moderately deep, and sandy loam to silty loam in texture [59,95,192]. Parent materials may include sandstone, limestone, pumice, and granite [25,53,95,138,214,230].
Detailed soil data for the Teakettle Experimental Forest in the Sierra Nevada  and a Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita community located in the Cascade Range of south-central Oregon  are available. Detailed descriptions of soil chemical and mineral composition in a midseral Jeffrey pine/greenleaf manzanita forest are also available .SUCCESSIONAL STATUS:
Effect on conifers: Greenleaf manzanita inhibits the growth of Pacific ponderosa pine (Pinus ponderosa var. ponderosa) [10,34,129,142,186,202,219] and coast Douglas-fir (Pseudotsuga menziesii var. menziesii) [101,204] seedlings. Greenleaf manzanita's ability to severely deplete available soil moisture is the greatest contributing factor in greenleaf manzanita inhibition of Pacific ponderosa pine seedling growth [172,173]. The ability to extract water from moisture-depleted soil is likely attributable to a better-developed root system than that of conifer seedlings.
In plots with 0% greenleaf manzanita cover and 100% Pacific ponderosa pine seedling cover, Pacific ponderosa pine seedling production 2.5 years after establishment averaged 5 kg/300 m². In plots with 25% greenleaf manzanita and 75% Pacific ponderosa pine cover, seedling production was reduced to 2 kg/300 m², a 60% decrease. Anderson and Helms  found that greenleaf manzanita seedling mean root:shoot ratio and mean root:total root length were 30% and 35%, respectively: greater than ponderosa pine seedling ratios. Tinnin and Kirkpatrick  found that Douglas-fir seedling root growth was significantly (P<0.05 and P<0.001) less when growing in greenleaf manzanita leaf litter than in the absence of greenleaf manzanita leaf litter. In a review by Tappeiner and others , greenleaf manzanita was described as reducing pine (Pinus spp.) volume by 9% to 37%, reducing conifer height, crown volume, and biomass, and causing ponderosa pine moisture stress that decreased productivity.
In a review by Aune , increased shrub cover (greenleaf manzanita, snowbrush ceanothus, and Klamath plum (Prunus subcordata)) is shown to decrease Pacific ponderosa pine survival and growth rates.
|Effects of shrub interference on survival,
height, and diameter of Pacific ponderosa pine
at Mt Shasta, California, over an 18-year period
|Pacific ponderosa pine characteristics||% shrub cover|
Greenleaf manzanita fields may cause substantial mortality of white fir and sugar pine seedlings. On the Teakettle Experimental Forest east of Fresno, California, fields of greenleaf manzanita were burned in October 1999, either lightly (no added fuel, some foliage burned) or with a "hot" fire (fuels added before burn, near complete foliage burned off and some stems charred). Burn and unburned control plots were planted with either white fir or sugar pine seedlings in October 2000. At the end of the second growing season (October 2002), total white fir survival rate across both treatments was 0.6%, which was significantly lower (P=0.02) than total survival of sugar pine (3.8%). Only early on (spring 2001) was there a significant difference (P=0.01) between treatments and survival rates. In spring 2001 white fir and sugar pine survival was lower on "hot" burn sites for both species than the light-fuel treatments .
Fire: Within its range, greenleaf manzanita is very prevalent as a pioneer species following fire [50,66,123,125,181,229]. Following crown fires in white fir and yellow pine forests of the Sierra Nevada, burn sites are "rapidly" occupied by montane chaparral shrub species including greenleaf manzanita [44,97,199].
In the pinyon-juniper (Pinus monophylla, P. edulis-Juniperus occidentalis) woodlands of the San Bernardino Mountains at elevations greater than 7,000 feet (2,000 m), greenleaf manzanita had 10.7% cover and a density of 1,416/ha on 70-year-old burn sites. Following fires in the pinyon-juniper woodlands, which were mostly severe canopy fires, shrub cover and density increased for 30 to 50 years. Based on observation from other sites and aerial photography, chronosequence sampling, and replication of the 1929 to 1935 California Vegetation Type Map, the researchers concluded that mature shrubs acted as nurse plants for singleleaf pinyon, which established beneath shrubs at approximately 25 to 40 years after fire. Around postfire year 50, shrubs began to decline, and by postfire years 100 to 150, a mature woodland returned .
Logging: Greenleaf manzanita thrives following logging in mixed-conifer forests [61,72,228]. Forty years following logging of mixed-conifer forests on the Blacks Mountain in northeastern California, greenleaf manzanita cover was significantly greater (P<0.05) on 30-year-old cut sites than uncut sites [215,216].
|Greenleaf manzanita flowering period|
|Arizona||May to June |
|California||April to June |
|Nevada||April to June [106,192]|
|southern California||May to June |
|Utah||April to June |
|Great Basin||May to June |
|Pacific Northwest||May to June |
|Baja California||March to June |
Flowering of greenleaf manzanita may be triggered by summer moisture stress . The number of flowers produced is partially dependent upon the amount of the previous year's precipitation. Flower buds form 1 year prior to maturity. They are dormant the following summer, fall, and winter, and bloom the next spring .
Fruits ripen in late summer to early fall . Generally, this species fruits over its entire range between July and October [20,217]. In Nevada, fruiting occurs from May to September . The fruits may occasionally persist on the shrub year-round . Most chaparral species experience the greatest amount of growth in May and June. Growth ceases in mid-July due to high air temperatures and low soil moisture .
The seasonal development of greenleaf manzanita was as follows in a Pacific ponderosa pine forest :
|Greenleaf manzanita phenological development in central Oregon|
|6 May||Few blossoms beginning to develop|
|27 May||Blossoming near completion|
|5 July||Blossoming complete and new fruits formed|
|22 September||No visible sign of growth|
Fire regimes: Greenleaf manzanita is found in plant communities/ecosystems with varying fire regimes. Greenleaf manzanita may occur where fire-return intervals are as short as 1 year or as long as several hundred years. A more detailed description of greenleaf manzanita fire-return intervals follows.
Oak/chaparral woodland: The oak/chaparral woodland of Yosemite National Park, California, had a presettlement fire-return interval of 20 to 30 years. Presently, many areas have not burned for 60 to 100 years due to fire exclusion .
Lodgepole pine forests: Greenleaf manzanita occurs in lodgepole pine forests on the east side of the Cascade Range, the Blue Mountains, and the Okanogan Highlands of Washington north into British Columbia and south to Colorado and California. Inland Pacific Northwest lodgepole pine communities had an historic mean fire-return interval of 112 years. Lodgepole pine forests in areas susceptible to summer drought in the Inland Pacific Northwest historically had low- to medium-severity surface fires occurring at intervals of 25 to 50 years .
Mixed-conifer forests: Greenleaf manzanita is prevalent in open mixed-conifer forests, particularly in the Sierra Nevada. The minimum and maximum historic fire-return intervals for mixed-conifer forests of the Sierra Nevada ranged from approximately 3 to 8 years and 10 to 20 years, respectively [44,107,126,155,165]. Prior to fire exclusion in the Sierra Nevada, fires were generally low -severity surface fires, and fire-return intervals were frequent .
Elsewhere, the fire-return interval ranges from 15 to 29 years in mixed-conifer/chaparral sites in the San Bernardino Mountains , from 10 to 80 years in mixed conifer-hardwood forests of southwestern Oregon , and 4 to 7 years in mixed-conifer forests in Bryce Canyon National Park, Utah .
Using aerial photography and ground sampling, the approximate mean fire-return interval from 1925 to 1991 for the Sierra San Pedro Mártir, Baja California, was estimated at 52 years. The Sierra San Pedro Mártir is the last remaining mixed-conifer forest along the Pacific coast still subject to uncontrolled, periodic surface fires. It is estimated that the infrequent fires are severe surface fires. Heavy fuels result from the gradual buildup of shrub cover (predominately greenleaf manzanita, pointleaf manzanita (Arctostaphylos pungens), and pinkbracted manzanita (A. pringlei ssp. drupacea)), conifer recruitment, and litter accumulation .
Montane chaparral: Fire-return intervals for montane chaparral sites in the Sierra Nevada are variable . Nagel and Taylor  studied the fire ecology of 6 montane chaparral sites codominated by greenleaf manzanita and huckleberry oak (Quercus vaccinifolia) in Lake Tahoe Basin in the northern Sierra Nevada. Prior to fire exclusion in the late 1800s and early 1900s, the mean fire-return interval was 28 years, with a range of 16 to 40 years. Other research estimates that the fire-return interval ranged from 30 to 60 years . When fuel moistures drop below 100%, fire is likely to remove a large portion of the shrubs in this type. When moisture drops below 100%, explosive fire conditions with rapid rates of spread and flame heights greater than 10 feet (3 m) are expected . High-severity fires are typical, with most shrubs being top-killed .
Pinyon-juniper woodlands: In the San Bernardino Mountains, greenleaf manzanita occurs in high-elevation (7,000 feet (2,000 m)) pinyon-juniper woodlands. Using fire scar data, Wangler and Minnich  estimate the fire-return interval was approximately 480 years. When fire occurs, it is generally high-severity crown fire .
Pacific ponderosa pine forests: Fire scar data from the years 1700 to 1875 on dry ridges in the southern Sierra Nevada showed that the minimum and maximum fire-return intervals were 2 and 12 years, respectively. The mean fire-return interval for that period was 5.5 years . Fire historically occurred approximately every 13 years in ponderosa pine woodlands of Lava Bed National Monument, California . Greenleaf manzanita occurs throughout ponderosa pine forests on the eastern side of the Oregon Cascade Range. Using fire scar analysis, Bork  found that the mean historic fire-return intervals at 3 sites, Cabin Lake, Pringle Butte, and Lookout Mountain, were 24, 11, and 15 years, respectively. In a Pacific ponderosa pine/incense-cedar (Calocedrus decurrens)/greenleaf manzanita community, the fire-return interval ranged from 9 to 42 years .
Red fir forests: For the period 1740 to 1985, the mean fire-return interval at Swain Mountain Experimental Forest in northeastern California was 12.9 years, with a range of 1 to 57 years. Larger fires (>10 acres (5 ha)) occurred every 26.2 years, with a range of 11 to 47 years. Average fire-free intervals were shorter during the settlement period (1851-1934, x = 7.9 years) than during the presettlement (1740-1850, x = 21.4 years) and fire-exclusion (1935-1985, x = 17.3 years) periods. Fires in red fir forests are generally of low and moderate severity .
The following table provides fire-return intervals for plant communities and ecosystems where greenleaf manzanita is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
|Fire-return intervals for plant communities with greenleaf manzanita|
|Community or Ecosystem||Dominant Species||Fire Return Interval Range (years)|
|California chaparral||Adenostoma and/or Arctostaphylos spp.||<35 to <100|
|California montane chaparral||Ceanothus and/or Arctostaphylos spp.||50-100 |
|sugarberry-America elm-green ash||Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica||<35 to 200 |
|paloverde-cactus shrub||Cercidium spp./Opuntia spp.||<35 to <100 |
|curlleaf mountain-mahogany*||Cercocarpus ledifolius||13-1,000 [8,184]|
|western juniper||Juniperus occidentalis||20-70|
|Rocky Mountain juniper||Juniperus scopulorum||<35|
|pinyon-juniper||Pinus-Juniperus spp.||<35 |
|Rocky Mountain lodgepole pine*||Pinus contorta var. latifolia||25-340 [16,17,201]|
|Sierra lodgepole pine*||Pinus contorta var. murrayana||35-200 |
|Colorado pinyon||Pinus edulis||10-400+ [64,71,115,168]|
|Jeffrey pine||Pinus jeffreyi||5-30|
|Pacific ponderosa pine*||Pinus ponderosa var. ponderosa||1-47 |
|interior ponderosa pine*||Pinus ponderosa var. scopulorum||2-30 [5,13,132]|
|Arizona pine||Pinus ponderosa var. arizonica||2-15 [13,48,185]|
|quaking aspen (west of the Great Plains)||Populus tremuloides||7-120 [5,81,151]|
|Rocky Mountain Douglas-fir*||Pseudotsuga menziesii var. glauca||25-100 [5,6,7]|
|coastal Douglas-fir*||Pseudotsuga menziesii var. menziesii||40-240 [5,156,178]|
|Pacific coast mixed evergreen||Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii||<35-130 [5,39]|
|California oakwoods||Quercus spp.||<35 |
|oak-juniper woodland (Southwest)||Quercus-Juniperus spp.||<35 to <200 |
|coast live oak||Quercus agrifolia||2-75 |
|canyon live oak||Quercus chrysolepis||<35 to 200|
|Oregon white oak||Quercus garryana||<35 |
|California black oak||Quercus kelloggii||5-30 |
|redwood||Sequoia sempervirens||5-200 [5,63,195]|
|Greenleaf manzanita seedlings in postfire year 1. Sequoia-Kings National Park, California. Photo courtesy of Eric Knapp.|
Greenleaf manzanita with lignotubers sprout after fire unless the entire periphery of the lignotuber is deeply charred, which rarely happens [15,181]. Shrubs produce new sprouts from dormant buds on the lignotuber in as little as 10 days to 3 weeks [31,93,99,157]. Greenleaf manzanita plants with a lignotuber can withstand repeated burnings [31,52]. However, Martin  found that repeated burning of greenleaf manzanita plants with a lignotuber can cause plant mortality.DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Prescribed burning increased greenleaf manzanita density on the Deschutes National Forest. During the spring of 1979, moderate- and high-severity prescription burns were conducted in a Pacific ponderosa pine/antelope bitterbrush-greenleaf manzanita/western needlegrass community. Prior to burning, 300 to 400 greenleaf manzanita shrubs/ha occurred on both treatment sites. In postfire year 15 (1984), greenleaf manzanita averaged 2,590 shrubs/ha on moderate-severity burns and 3,550 shrubs/ha on high-severity burn sites .
On Donner Pass, California, an August 1960 wildfire burned 40,000 acres (16,000 ha) in a conifer forest consisting primarily of mixed pine and fir species. In 1963 and 1975 line transects were run throughout the burned areas. In 1963 greenleaf manzanita was 0% on line transects. Greenleaf manzanita cover on transects increased to 3.8% in 1975 .
Seedlings: In the following studies, an increase in greenleaf manzanita seedlings occurred following fire. Higher-severity fires caused greater seedling establishment than lower-severity fires [109,111,112,222].
Before and after a September 1927 fire on a chamise-dominated site in Mendocino County, California, greenleaf manzanita seedlings/milacre were counted. Prior to the fire, there were no greenleaf manzanita seedlings present. Seedling establishment was greatest during the first postfire year. A survey of the area during postfire year 7 (1934) found that a large proportion of greenleaf manzanita seedlings which were present in postfire year 5 had survived and developed into robust plants .
|Greenleaf manzanita seedlings/milacre before and 1 to 5 years after burning|
|Prefire||1st postfire year||2nd postfire year||3rd postfire year||4th postfire year||5th postfire year|
Sixty to 70-year-old white fir stands with an understory of snowbrush ceanothus and greenleaf manzanita and/or pinemat manzanita (species not differentiated in study) were burned before and after tree harvest in the Lassen National Forest, California . Prior to spring or fall burning and white fir harvest in 1983, manzanita seedlings ranged from 0 to 1,700 seedlings/acre. During the summer of 1984, manzanita seedling density ranged from 11,000 to 16,000/acre on sites burned in spring 1983 before harvest. Manzanita seedling densities on sites burned after harvest in the spring and fall of 1983 ranged from 13,000 to 16,000/acre and 10,000 to 11,000/acre for spring and fall burns, respectively. The density of manzanita seedlings after postharvest burning was considerably higher for spring burns than for fall burns. This seasonal difference was attributed to consistently higher percent burned area in spring than in fall; fall fuels were wet due to rains .
Fall and spring prescribed burns of varying severities conducted in 1983 at 2 sites on the west slope of the Sierra Nevada and Cascade Range produced significant greenleaf manzanita seedling increases (P<0.10) during either postfire year 1 or 2 [109,111,112]. The first site, Blodgett Forest Research Station, sits at an elevation of 4,300 feet (1,300 m) and was dominated by white fir, ponderosa pine, incense-cedar, and Douglas-fir, with a scattered and diverse shrub understory consisting of whitethorn ceanothus, greenleaf manzanita, California black oak, deer brush (Ceanothus integerrimus), giant chinquapin (Chrysolepis chrysophylla), tanoak (Lithocarpus densiflora), Sierra mountain misery (Chamaebatia foliolosa), and pine rose (Rosa pinetorium). The second site, Quincy Ranger District, sits at an elevation of 4,432 feet (1,351 m) and was dominated by Jeffrey pine, coast Douglas-fir, and incense-cedar. The most common understory shrubs included California black oak, deer brush, thimbleberry (Rubus parviflorus), and pale serviceberry (Amelanchier pallida). Greenleaf manzanita was a minor species at Quincy; therefore, greenleaf manzanita seedlings were far fewer after Quincy fires than fires at Blodgett [109,111,112].
Four burn treatments were conducted at each site. Researchers included an early spring-moderate consumption burn; a late spring-high consumption burn; an early fall-high consumption burn; and a late fall-moderate consumption burn. The early spring-moderate consumption burn was conducted as soon as fuels could carry fire. The late spring-high consumption burn was conducted as late in the season as safely possible (prior to the active fire suppression season). The early fall-high consumption burn was conducted as early in the fall as possible and before major precipitation. Finally, the late fall-moderate consumption burn was conducted 1 to 2 days after significant precipitation. Seedling densities of greenleaf manzanita, particularly at the Blodgett site, were much higher on high-consumption burns than moderate-consumption burns. Comparing high-consumption burns, spring burns produced more greenleaf manzanita seedlings than fall burns at Blodgett, while the opposite was true at Quincy. On spring sites, excluding the early spring-moderate consumption burn at the Blodgett site, greenleaf manzanita seedlings were not produced in postfire year 1 because the stratification period was incomplete until the winter of 1985. At 1 Blodgett site (late fall-moderate consumption) and 2 Quincy sites (early fall-high consumption and late fall-moderate consumption), greenleaf manzanita seedlings increased in postfire year 1 but decreased during postfire year 2. This was attributed to shading: greenleaf manzanita seedlings occurred under a dense canopy [109,111,112].
Most greenleaf manzanita seeds present in the seed bank had been cached by rodents many years prior to the fires, likely before conifer establishment. After postfire seedling establishment, the rodents dug into these caches in search of seeds. This caused substantial mortality of fragile greenleaf manzanita seedlings .
The table below describes density of greenleaf manzanita seedlings prior to the prescription burns (1983), at the end of the 1984 and 1985 growing seasons (postfire years 1 and 2), and on the unburned control sites [109,111,112].
|Mean (SE) density/ha of greenleaf manzanita seedlings at the Blodgett site|
|Treatment||Date of fire||1983 (prefire)||1984||1985|
|Early fall-high consumption||28 Sept. 1984||0 (0)||ND*||11,833 (2,625)|
|Late fall-moderate consumption||8 Oct. 1983||2,042 (1,116)||4,875 (1,508)||958 (305)|
|Late spring-high consumption||26 June 1984||367 (200)||0 (0)||25,233 (5,832)|
|Early spring-moderate consumption||17 May 1984||233 (202)||34 (33)||4,600 (2,059)|
|Control||467 (210)||134 (65)||134 (65)|
|Mean (SE) density/ha of greenleaf manzanita seedlings at the Quincy site|
|Treatment||Date of fire||1983 (prefire)||1984||1985|
|Early fall-high consumption||15 Sept. 1983||0 (0)||3,567 (769)||861.1 (285)|
|Late fall-moderate consumption||12 Oct. 1983||0 (0)||1,167 (356)||167 (87)|
|Late spring-high consumption||24 May 1984||0 (0)||0 (0)||833 (540)|
|Early spring-moderate consumption||7 May 1984||0 (0)||0 (0)||367 (155)|
|Control||0 (0)||0 (0)||0 (0)|
For a complete review of this study, see the Research Project Summary by Kauffman [109,111,112].
Repeated burning: On Lookout Mountain in central Oregon, a Pacific ponderosa pine/greenleaf manzanita community was burned twice over a 4-year period, causing nearly complete mortality of mature greenleaf manzanita and complete mortality of greenleaf manzanita seedlings. Prescription burning took place at 2 sites, designated as the upper and lower plots .
|Date, fuel, and weather conditions during greenleaf manzanita prescribed burns|
|Date/conditions||Upper plot, 1st burn||Lower plot, 1st burn||Upper plot, 2nd burn||Upper plot, 2nd burn|
|Relative humidity (%)||37-61||30-38||38-54||42-43|
|Fuel moisture content/new litter (%)||12-15||10-11||7-20||5-8|
|Fuel moisture content/old litter (%)||10-36||11-15||11-21||6-14|
|Fuel moisture content/duff (%)||26-65||26-75||14-21||52-54|
Following the fires, Martin  measured the percent of greenleaf manzanita that was unburned, burned and sprouting after fire, or killed. Greenleaf manzanita seedling mortality was measured after second fire. Unfortunately, the researcher did not provide the date of postfire vegetation measurements. "Poor" postfire sprouting of greenleaf manzanita occurred on both sites. The researcher expected this, since greenleaf manzanita is a poor sprouter in the area and the overstory was a closed Pacific ponderosa pine canopy. Results of the 2 fires are presented below :
Cover (%) of greenleaf manzanita
|Fire effects||Upper plot||Lower plot|
|Shrub seedlings dead||100||100|
For further information on prescribed fire use and postfire responses of multiple plant species in this plant community, see Martin's  original Research Paper.
In a montane chaparral site in northern California, a prescribed fire caused the germination of thousands of greenleaf manzanita seedlings. Three years later the site was reburned, and only 1 greenleaf manzanita seedling emerged after fire. A third and fourth fire completely eliminated greenleaf manzanita . Presumably, the initial fire exhausted the greenleaf manzanita seed bank, leaving few or no seeds to germinate following subsequent fires. Since it can take 8 to 10 years for greenleaf manzanita plants to flower and fruit , there was no replenishing of the seed bank between the first and second fires.FIRE MANAGEMENT CONSIDERATIONS:
Fire behavior: The fire behavior of 16 prescribed burned plots in montane chaparral communities codominated by greenleaf manzanita, huckleberry oak, and whitethorn ceanothus was measured in Yosemite National Park. These fires were separated in space and time. At the time of burning few shrubs exceeded 6 feet (2 m) in height .
|Fire behavior in a montane chaparral site in Yosemite National Park|
|Spread/minute (feet)||Flame length (feet)|
Fire economics: Literature is available on the costs associated with and the economic feasibility of burning greenleaf manzanita and other northern California chaparral species .
Live fuel moisture: In the Sierra Nevada and Cascade Range of California, greenleaf manzanita live fuel moisture during the 1984 fire season ranged from a high of approximately 150% on 29 May to a low of approximately 100% on 15 October . On the Shasta-Trinity National Forest, greenleaf manzanita mature leaf moisture content varied widely throughout the year. Samples taken in November 1966 showed an average moisture content of 120%, which dropped to 86% by 25 June 1967. From the end of June to 10 July 1967, average moisture content increased rapidly to 125% and remained at or above 100% until April 1968. The moisture content of new leaves was substantially higher than mature leaves. On 13 July, when the moisture content of mature leaves was 123%, new greenleaf manzanita leaves had a moisture content of 259% .
Physical burning characteristics: On the eastern slopes of Mt Shasta at elevations ranging from 4,000 to 6,000 feet (1,200-1,800 m), greenleaf manzanita physical characteristics relevant to fire were studied over a 4-year period .
|Average greenleaf manzanita physical characteristics|
|Characteristics||Foliage||Woody fuel (inches diameter)|
|< 0.25||0.26-0.50||0.51-1.0||> 1.0|
|Living fuel density (lb/ft³)||54.7||38.9||41.3||44.5||46.4|
|Solvent extractives in living fuel (%)||16.9||6.2||4.2||3.6||3.4|
|Solvent extractives in dead fuel (%)||0||2.6||1.2||1.4||1.9|
|Surface-to-volume ratio (ft²/ft³)||1,623||343||137||66||32|
|Heating value of living fuel (Btu/lb)||9,076||8,246||8,212||8,306||8,320|
|Heating value of dead fuel (Btu/lb)||no data||8,280||8,367||8,201||8,429|
Data on the average greenleaf manzanita biomass by fuel size class and crown cover across many areas of Oregon, Washington, and northern California east of the Cascade Range are available .
Soil: Data on the average bulk density and nutrient concentrations of soil on burned shrub sites dominated by snowbrush ceanothus and containing greenleaf manzanita are available in Johnson and others .Surface and subsurface (3 to 4.3 inches (7-11 cm)) soil temperatures were measured during prescribed fires of high and low severity in greenleaf manzanita stands in the Sierra National Forest, California, at an elevation of 7,000 feet (2,000 m). High-severity fires were created by adding fine and woody fuels to the burn sites. Low-severity sites were burned with existing fuel loads. On the high-severity sites, temperatures reached over 600 °F (300 °C) at the soil surface and 400 °F (200 °C) in the subsurface zone. On the low-severity sites, temperatures reached 300 °F (150 °C) at the surface and 200 °F (100 °C) in the subsurface zone .
The fruits and seeds of greenleaf manzanita are important to a variety of birds and mammals [86,122,146,149]. The fruits are a food source for American black bears in northern California and southern Oregon during late summer and early fall [69,94,95,97,169]. Grouse, wild turkeys, songbirds, and deer mice also consume the fruits [85,94,97,100].
Palatability/nutritional value: The palatability of greenleaf manzanita is described as "poor to worthless" for domestic goats and "worthless" for cattle and horses [104,183]. It is palatable to domestic sheep [79,144] (see Other Management Considerations).
The crude protein of greenleaf manzanita taken from California ranges from a low of 5.2% in February to a high of 7.8% in August and September . In Truckee River Canyon on the California-Nevada border, crude protein of greenleaf manzanita ranged from a low of 6.0% in January to 7.8% in August and September . The ash content of greenleaf manzanita foliage taken from Mt Shasta at elevations of 4,000 to 6,000 feet (1,200-1,800 m) averaged 3.9% .
Cover value: While there is little in the literature on the cover value of greenleaf manzanita, it is likely valuable to numerous animal species due to the dense thickets it produces. It is described as an important cover species for small mammals, birds, insects, and arthropods [33,94,208].VALUE FOR REHABILITATION OF DISTURBED SITES:
A greenleaf manzanita cultivar ('Altura') is available .OTHER USES:
Greenleaf manzanita had many uses for Native Americans. The fruits were eaten whole, made into cider and jelly, and brewed into tea to treat poison-oak (Toxicodendron diversilobum) exposure [94,200]. The leaves were used as an emetic, for treating insect bites, and to relieve bronchitis, dropsy, and other diseases [94,200]. Dried leaves were also used in herbal smoking mixtures [59,200].OTHER MANAGEMENT CONSIDERATIONS:
Herbicides: Research papers are available on the effects of greenleaf manzanita herbicide treatments on conifer species [40,41,45,75,121,140,143,147,148,221].
Host species: Greenleaf manzanita is a host to the manzanita leaf-gall aphid, which produces galls on the leaves and flower buds . On the Deschutes National Forest, greenleaf manzanita is host to at least 12 fungal species, 3 of which are "important" plant pathogens .Mechanical control: While the work is arduous, greenleaf manzanita can be successfully controlled by grubbing. In a young northern California Pacific ponderosa pine plantation, greenleaf manzanita was significantly reduced (P<0.05) over a 10-year period on grubbed sites . Other mechanical control studies are available .
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